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Arquivos do Museu Nacional, Rio de Janeiro, v.65, n.4, p.417-459, out./dez.2007 ISSN 0365-4508

THE FIRST “PROTOSUCHIAN” (ARCHOSAURIA: ) FROM THE () OF 1 (With 16 figures)

LUCAS E. FIORELLI 2 JORGE O. CALVO 3

ABSTRACT: The remains of “protosuchians” from the Cretaceous come, to exception of “Las Hoyas crocodyliform” from the Lower Cretaceous of Spain, exclusively of Central : Zaraasuchus, , Zosuchus, and Artzosuchus from the Upper Cretaceous of Mongolia; Tagarosuchus from Lower Cretaceous of Southern Siberia; , , and Shantungosuchus from Lower Cretaceous of . We report a new basal crocodyliform taxon, Neuquensuchus universitas gen.nov., sp.nov., from Neuquén Province, , belonging to Bajo de la Carpa Formation, representing the first and only “protosuchian” from the Cretaceous of Gondwana. The articulated and fragmentary materials belonged to a willowy, slender species, with very long and thin extremities. As in Shantungosuchus, the cervical centers are lengthened, with prominent ventral keel and well developed anteroventral parapophyses. As in basal crocodylomorphs, it possesses two sacral vertebrae. Also, a much enlarged scapular blade, with well developed acromial ridge and the posterior edge similar to Sichuanosuchus. The pronounced deltopectoral crest in the complete is equivalent to Sichuanosuchus and as this, a circular, elongated and thin shaft with the medial condyle longer than the lateral one. Also, the complete and is similar in their proportions to Sichuanosuchus. As this, the pubis is lengthened, very thin in the half section and not very expanded distally. The femur, tibia and fibula are elongated and similar to other non-derivated crocodyliforms. Besides representing the first Cretaceous “protosuchian” of Gondwana, the occurrence of these outside of Asia and during the Cretaceous offers new evidence of pre- dispersion between Gondwana and Central Asia through Europe. Key words: . Protosuchian. Neuquensuchus universitas gen.nov., sp.nov. Cretaceous. Gondwana. RESUMEN: El primer “protosuquio” (Archosauria: Crocodyliformes) del Cretácico (Santoniano) de Gondwana. Los restos de “protosuquios” del Cretácico provienen, a excepción del “crocodyliforme de Las Hoyas” del Cretácico Inferior de España, exclusivamente de Asia Central: Zaraasuchus, Gobiosuchus, Zosuchus y Artzosuchus del Cretácico Superior de Mongolia; Tagarosuchus del Cretácico Inferior del sur de Siberia; Edentosuchus, Sichuanosuchus y Shantungosuchus del Cretácico Inferior de China. Aquí reportamos un nuevo taxón de crocodyliforme basal, Neuquensuchus universitas gen.nov., sp.nov., de la provincia de Neuquén, Argentina, correspondiente a la Formación Bajo de la Carpa, representando el primer y único “protosuquio” del Cretácico de Gondwana. Los materiales fragmentarios y articulados corresponden a una especie esbelta y delgada, con extremidades largas y delgadas. Al igual que en Shantungosuchus, los centros cervicales son alargados, con una quilla ventral prominente y parapófisis anteroventrales bien desarrolladas. Como en los crocodyliformes basales, Neuquensuchus posee dos vértebras sacras. Además, una hoja escapular muy expandida, con un puente acromial bien desarrollado y el borde posterior similar a Sichuanosuchus. La cresta deltopectoral pronunciada en el húmero es equivalente a la de Sichuanosuchus y al igual que este, la diáfisis es circular, alargada y delgada con el cóndilo medial mayor que el lateral. Asimismo, las proporciones del radio y la úlna son similares a Sichuanosuchus. Como este, el pubis es alargado, muy delgado en su sección media y poco expandido distalmente. El fémur, tibia y fíbula son alargados y similares a otros crocodyliformes no derivados. Además de representar el primer “protosuquio” cretácico de Gondwana, su presencia fuera de Asia y Europa durante el Cretácico ofrece nueva evidencia de un evento de dispersión pre-Albiano entre Gondwana y Asia Central a través de Europa. Palabras clave: Crocodylomorpha. Protosuquio. Neuquensuchus universitas gen.nov., sp.nov. Cretácico. Gondwana.

1 Submitted on September 14, 2006. Accepted on October 24, 2007. 2 Centro Regional de Investigaciones Científicas y Transferencia Tecnológica (CRILAR). Entre Ríos y Mendoza s/n, CP 5301, Anillaco, La Rioja, Argentina. E-mail: [email protected]. 3 Centro Paleontológico Lago Barreales (CePaLB), Universidad Nacional del Comahue. Ruta Provincial 51, km 65, Neuquén, Argentina. 418 L.E.FIORELLI & J.O.CALVO

INTRODUCTION MATERIAL AND METHODS

Fossil remains of basal non- The remains were found and gathered by Mr. Oscar Crocodyliformes from Cretaceous come almost de Ferrariis (at that time Director of the Museum exclusively from the Asian continent, to exception of the National University of Comahue), together of “Las Hoyas Crocodyliform” (SANZ et al., 1988) with J.O.C. The materials of this new basal (Fig.1) from the Lower Cretaceous of Las Hoyas, crocodyliform were originally referred as Spain (upper ; DIÉGUEZ et al., 1995). (MUCPv-137) and were collected in The Asian forms are represented by species 1987. The study of the museum collection allowed coming from China, Mongolia and Russia. From us to find one more specimen represented by China comes Edentosuchus tienshanensis (YOUNG, fragmentary postcranial material but in good 1973; POL et al., 2004), a from the preservation (Fig.3). Lower Cretaceous of , Xinjiang; Institutional abbreviations: GMPKU, Geological Shantungosuchus hangjinensis (WU et al., 1994) Museum, School of Earth and Space Sciences, Peking from the Luohandong Formation, Zhidan Group, University, Beijing, People’s Republic of China; IGM, Inner Mongolia and Sichuanosuchus shuhanensis Mongolian Institute of Geology, Ulaan Bataar, (WU et al., 1997) from an uncertain locality of Mongolia; IVPP, Institute of Vertebrate Sichuan. From Mongolia come forms belonging and Paleoanthropology, Beijing, People’s Republic of to the age. Gobiosuchus kielanae China; LACM, Natural History Museum of Los Angeles (OSMÓLSKA, 1972; OSMÓLSKA et al., 1997) comes County, Los Angeles, California, USA; MACN, Museo from the Bayan Zak locality; Gobiosuchus Argentino de Ciencias Naturales, Buenos Aires, (?)parvus (EFIMOV, 1983), later considered Argentina; MUCP, Museo de Geología y Paleontología, conspecific of G. kielenae (OSMÓLSKA et al., 1997), Universidad Nacional del Comahue, Neuquén, comes from Üüden Sair locality; Zosuchus Argentina; UNC, Department of Geological Sciences, davidsoni (POL & NORELL, 2004a) and Zaraasuchus University of North Carolina at Chapel Hill; ZDM, shepardi (POL & NORELL, 2004b) come from Zos Zigong Museum, Zigong, Sichuan, China; Canyon locality; Artzosuchus brachicephalus ZPAL, Instytut Paleobiologii PAN, Warszawa, Poland. (EFIMOV, 1983), a very fragmentary form of uncertain filiation, comes from the same locality that G. (?)parvus. Lastly, Tagarosuchus kulemzini (ALIFANOV et al., 1999), with practically complete , comes from the Lower Cretaceous of Shestakovo locality, South Siberia. Here we present a new basal form of crocodyliform from the Upper Cretaceous of Northern Patagonia, Neuquén Province, Argentina. The remains come from the Bajo de la Carpa Formation, Neuquén Group (Fig.2), and represent the first “protosuchian” form for the Cretaceous of Gondwana. In this paper, we describe the anatomy of this new Crocodyliform together with a parsimony analysis of their Fig.1- Map of Eurasia showing the places of origin of the species of Cretaceous phylogenetic relationships. protosuchians.

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Fig.2- Up right: satellital map showing the location of Argentina and Patagonia in ; up left: satelital map of Northpatagonic region, showing the location of the Neuquén Province; below left: area of Comahue where were found and collected the materials of Neuquensuchus universitas, gen.nov., sp.nov. (scale bar = 10km - right inferior bar). Below right: stratigraphy of the Cretaceous of Neuquén Basin and stratigraphic column of the Neuquén Group (based on LEANZA et al., 2004. (Satellital images taken from GoogleEarth).

RESULTS dinosaur Alvarezsaurus calvoi BONAPARTE, 1991 and Velocisaurus unicus BONAPARTE, 1991; GEOLOGY sauropod as cf. Laplatasaurus (LEANZA et al., 2004), Titanosauridae indet. (CHIAPPE & The Río Colorado subgroup constitutes the top of CALVO, 1994; pers.obs.), Neuquensaurus sp. the Neuquén Group; it is widely distributed in the (pers.obs.), Antarctosaurus and the peculiar South of the Neuquén Basin. The subgroup is beaked sauropod Bonitasaura salgadoi divided in two formations: Bajo de la Carpa (lower) APESTEGUÍA, 2004. Birds as Neuquenornis volans and Anacleto (upper) (LEANZA et al., 2004) (Fig.2). Bajo C HIAPPE & C ALVO, 1994 and Patagopteryx de la Carpa Formation is composed of coarse- deferrariisi ALVARENGA & BONAPARTE, 1992, snakes grained, light violet and pink sandstones of fluvial as Dinilysia patagonica WOODWARD, 1901, bird origin. The age has been dated as Santonian (LEANZA eggs in nests (SCHWEITZER et al., 2002), dinosaur et al., 2004) (Fig.2). eggs named Megaloolithus patagonicus CALVO et Outcrops in the area have given a wide variety al., 1997. Crocodyles are represented by of fauna such as carnotaurine abelisaurid Notosuchus terrestris WOODWARD, 1896, theropod (PORFIRI & CALVO, 2006) and the avian brachybuccalis BONAPARTE, 1991,

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Cynodontosuchus rothi WOODWARD, 1896, and materials were collected. postcranials articulated remains of a new Holotype – MUCPv-47 (Fig.3). Six , peirosaurian crocodyliform (FIORELLI et al., 2007). first four dorsal vertebrae, two sacral vertebrae and The remains of this new “protosuchian” have been first five caudal vertebrae. Posterior cervical ribs gathered on the South margin of Neuquén River and anterior dorsal ribs. Fragmentary right (North Neuquén City) increasing the number of , humerus, ulna and rights radius; left crocodyliforms found in the formation. scapula and humerus. Right pubis, fragment of right ischium, femur, tibia and right fibula; SYSTEMATIC PALEONTOLOGY fragment of the left ilium. Referred specimens – MUCPv-161 (Fig.3). Proximal Crocodylomorpha WALKER, 1970 end of left tibia, distal end of left fibula and left Crocodyliformes HAY, 1930 astragalus. (sensu BENTON & CLARK, 1988) Type locality – The remains were found in the North WHETSTONE & WHYBROW, 1983 of the Neuquén City on the campus of the Universidad Nacional del Comahue (National Neuquensuchus universitas, nov. gen. et nov. sp. University of Comahue), Neuquén Province, Argentina (Fig.2). Etymology – Generic name “Neuquén” in reference to the Neuquén City; “suchus”, Greek Type horizon – Bajo de la Carpa Formation, Río for crocodyle. Specific name “universitas” in Colorado Subgroup, Neuquén Group (Santonian; reference to the universitary campus, where the LEANZA et al., 2004) (Fig.2).

Fig.3- Neuquensuchus universitas gen.nov., sp.nov. Referred material. MUCPv-47 (holotype): A, B and E; MUCPv-161 (referred specimens): C and D. A, cervical vertebrae, first dorsal vertebrae, left scapula and left humerus. B, sacral and first caudal vertebrae and right pubis, ischium, femur, tibia and fibula. C, left tibia and fibula. D, left astragalus. E, right humerus, ulna, radius and radial. (Abbreviations in the Appendix IV).

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Diagnosis – Relatively small, thin and slender articulate cervical vertebrae with the first four crocodyliform, diagnosed by the following dorsal, two sacral vertebrae and relatively well combination of poscranial characters: lengthened preserved five anterior caudal vertebrae that are cervical vertebrae with low ventral keel, articulated to the sacral . parapophysis and diapophysis anteroposteriorly Regarding the cervical section (Fig.4), this specimen lengthened. Neural spines elongated in dorsal possesses a relatively long and thin neck, similar vertebrae, with their centra lengthened without to those other basal crocodylomorphs, as for ventral keel but with a very low anterior example (CRUSH, 1984) and hypapophysis. Two laterally enlarged sacral Gobiosuchus (OSMÓLSKA et al., 1997). On the cervical vertebrae. First caudal vertebra with a tenuous sequence, the first one, here considered the fourth, opisthocoelous and elongated anterior caudal is incomplete, preserving just the posterior portion vertebra, relatively low. Scapula with an of the centrum (Fig.4). All cervical vertebrae and important dorsal expansion and a good preserved dorsal are slightly amphicoelous. The development of the posterodorsal hook. Humerus long and thin cervical centra are parallelogram- with a good development of the lateroproximal shaped in lateral view, with an elevation of the expansion, long and thin diaphysis of the anterior face of the centrum, similarly to humerus with the medial condyle biggest than Terrestrisuchus (CRUSH, 1984), the lateral one. Very lengthened and thin ulna, elaphros (WU & CHATTERJEE, 1993), Zaraasuchus with olecranon process. Very thin and proximally (POL & NORELL, 2004b, IGM 100/1321), expanded radius. Thin and long pubis with a very Shantungosuchus (YOUNG, 1961, IVPP V2484; WU light distal expansion. Non-sigmoid and et al., 1994, IVPP V10097) and other cervicals of lengthened femur, smaller than the tibia. Crocodylia (ROMER, 1956; HOFFSTETTER & GASC, 1969). Neuquensuchus possess medially constricted, well marked cervical centra, similar to some basal DESCRIPTION AND COMPARISIONS crocodyliforms, such as Zaraasuchus (POL & NORELL, 2004b) and Shantungosuchus (YOUNG, 1961; WU et AXIAL SKELETON al., 1994) and different to other protosuchids and mesoeucrocodylians, as Edentosuchus (LI, The specimen MUCPv-47 of Neuquensuchus 1985) and (WU & SUES, 1996; FIORELLI, universitas possesses incomplete axial remains but 2005; POL, 2005), that possess short and compressed in good preservation state. It includes the last six cervical centra, without medial constriction.

Fig.4- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-47. Cervical vertebrae in left lateral view. (Abbreviations in the Appendix IV).

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This structure indicates wide lateral movements NORELL, 2004b). Laterally, in the base of the neural of the long neck in this basal patagonian spines, there is a cavity between the pre and crocodyliform. In another sense, each one of the postzygapophysis, nearly delimited by a small cervical centra possesses a long keel that runs developed suprapostzygapophyseal lamina (Fig.5A). anteroposteriorly in the whole ventral surface, Prezygapophysis and postzygapophysis, in dorsal forming deep furrows toward both sides of this and view are robust, laterally high and slightly curved ventrally to the parapophysis (Figs.5B, 5C, 5D). laterally. Prezygapophysis articulate facets are Even so, the ninth centrum also possesses a less dorsomedially directed and postzygapophysis marked and lower keel, with shallow lateral furrows articulate facets are lateroventrally directed, like in than those present in anterior cervicals. These keels Zaraasuchus. Ventrally, the prezygapophysis are similar to those observed in “protosuchians” possesses a well developed lamina posteroventrally and notosuchians, like in the of directed, that continues with the anterior border of Shantungosuchus hangjinensis (WU et al., 1994, diapophysis; it directs anterodorsally the IVPP V10097), in the cervical vertebrae of prezygapophysis base (Fig.5D). There is a very (COLBERT & MOOK, 1951), marked border, that extends toward posterior among Sichuanosuchus huidongensis (PENG, 1996), the articular facets of the pre and postzigapophysis, Notosuchus (POL, 2005; FIORELLI, 2005, MACN-RN on the whole lateral surface of the neural 1037 and MUCPv-137) and (WU pedicelous. Similar condition has been observed & SUES, 1996, p.692-693, IVPP V8274) but the long in Zaraasuchus (Fig.5D). extension is a plesiomorphic character. The Regarding the dorsal vertebrae, only the first four parapophysis are very wide, well developed and have been preserved, with their corresponding robust with a lengthened articulate facet for the articulate ribs (Fig.6). It is observed that these capitulum of the cervical ribs (Figs.5C, 5D). The dorsals, corresponding to the tenth to twelfth articulated facets of these parapophysis possess vertebrae, possess the same anteroposterior lenght, an antero-lateroventral direction, similar to other but they fall in relation with the posterior cervical basal crocodylomorphs as in the first cervical ones ones. In Notosuchus and other Metasuchia there is of Terrestrisuchus (CRUSH, 1984), in the posterior a light increase in the longitude of the tenth (last cervical vertebra of Zaraasuchus (POL & NORELL, cervical in Notosuchus) and eleventh dorsal centrum, 2004b) or in the axis of Shantungosuchus (YOUNG, compared with the short cervical ones (POL, 2005; 1961; WU et al., 1994). Lateroventrally projected FIORELLI, 2005). All the centra are amphicoelous and parapophysis of Neuquensuchus universitas strongly constrained in the half section. Therefore, possesses a long parapophyseal ridge posteriorly. proximal and distal facets are very wide and inflated Posterior cervical vertebrae have the surfaces for (Figs.6C, 6D) like in Sichuanosuchus huidongensis the capitulum enlarged and lengthen, covering (PENG, 1996). The first dorsal vertebra does not practically the anterior half of the extensive possess a ventral kill and a true reduced centrum (Fig.5D). Between the parapophysis and hypapophysis appears (Fig.6D). In the first two diapophysis there is a prolonged depression, this dorsal vertebrae, the parapophyses are anteriorly character has been recorded in Zaraasuchus (POL located, ventrally directed and rounded. The third & NORELL, 2004b) and Protosuchus (COLBERT & MOOK, dorsal vertebra has the parapophysis small and 1951). The diapophyses are lengthened in the first dorsoventrally longer. Diapophyses are well cervical vertebra and they are anteriorly located developed. The cavities in the base of the neural below the neurocentral sutures. Nevertheles, in the spines are wider and shallower, not very deep but seventh cervical, the diapophyses are limited posteriorly by high and well-developed anteroventrally located on the suture. In the eighth suprapostzygapophyseal laminae (Figs.6A, 6B). In cervical, the diapophysis spreads rounding the lateral view, neural spines in anterior dorsals are tubercular process. Lastly, in the ninth cervical, very elongated and laminar (Figs.6A, 6B). the diapophysis is located more dorsally, as in In MUCPv-47, the poorly preserved sacral Terrestrisuchus. All cervicals possess an important vertebrae are articulated with the anterior five postdiapophyseal ridge, like in Zaraasuchus. The caudals (Fig.7). They are jointed by a suture. neural spines are not complete but they seem to Centra are short and very wide, flat and massive. be high and dorsoventrally lengthened, centrally (Figs.7C, 7E). The preserved transverse processes located in the neural arches, contrary to the seem to have been wide, similar to those of basal posterior cervical vertebrae of Zaraasuchus (POL & crocodylomorphs as (SUES et al., 2003).

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Fig.5- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-47. Posterior cervical vertebrae. A, right lateral view; B and D, left lateral view; C, ventral view. (Abbreviations in the Appendix IV).

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Anterior caudals correspond to the five firsts prezygapophysis, in the third and fourth caudals, (Figs.7A, 7B). Just centra and some pre and are inclined ventromedially. Hemals arches have postzygapophyses are preserved; they are more not been preserved but the articulated surfaces rounded and lengthened than in Notosuchus. In for the same one appear from the second caudal Neuquensuchus universitas centra are similar to vertebra. the first caudal vertebrae of Shantungosuchus (WU

et al., 1994) and other basal crocodyliforms. The APPENDICULAR SKELETON first caudal possesses a centrum very slightly opisthocoelic. Transverse processes in the second MUCPv-47 includes both scapulae, the left and third caudals are slightly square in humerus (Fig.6), ulna and right radius, left transverse section and they placed at the same ilium, right pubis, proximal right ischium, level than the zygapophysis. Pre and femur, tibia and fragment of the right fibula. postzygapophyses, in caudals, do not possess an MUCPv-161 includes a very well preserved extensive dorsal development as those in proximal left tibia, distal left fibula, and Notosuchus and other notosuchian and fragmentary remains of tarsus – left astragalus neosuchian, such as in (Fig.3). It is referred to Neuquensuchus due to (BUCKLEY & BROCHU, 1999) and their characters and similar proportions with (SCHWARZ et al., 2006). Articulation surfaces of the MUCPv-47.

Fig.6- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-47. First dorsal vertebrae. A and B, right lateral view; C and D, ventral view. (Abbreviations in the Appendix IV).

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SCAPULA Another important characteristic is the relationship between the dorsoventral length of The scapula of Neuquensuchus universitas (Figs.6, scapula and the total length of the humerus; only 8) is quite similar to that of Notosuchus (POL, 2005; in Terrestrisuchus, Gobiosuchus, Sichuanosuchus FIORELLI, 2005) and Sichuanosuchus shuhanensis (WU and Neuquensuchus universitas this scapular et al., 1997, IVPP V12088). However it differs from longitude represents less than 70% of the Notosuchus in having a less marked constriction longitude of the humerus, while in the remaining above the ventral expansion and a slender dorsal crocodylomorphs – included all the Metasuchia –, expansion. In notosuchians the dorsal expansion is it is always bigger. very developed and more anteroposteriorly extensive (POL, 2005; FIORELLI, 2005). As in S. shuhanensis, HUMERUS Neuquensuchus universitas has the anterior concave border of the scapular blade wider than the posterior MUCPv-47 preserves both humera (Figs.9-10). one and a well-developed acromial ridge, extended They are very long and thin (100.8mm), and along the anterior margin of ventral portion (Fig.8). similar in all its proportions and characteristic The hook, or projection in the posterodorsal vertex, to that of Gobiosuchus kielanae (OSMÓLSKA et al., is posteriorly directed and the dorsal border is 1997, ZPAL MgR-II/67), Zaraasuchus (POL & convex. It is only shared with Sichuanosuchus N ORELL, 2004b, IGM 100/1321), and shuhanensis (WU et al., 1997) and also in part with Sichuanosuchus shuhanensis (WU et al., 1997, Sichuanosuchus huidongensis (PENG, 1996). The hook IVPP V12088). The relationship between the is also visible in some sphenosuchians as in distal extension of the deltopectoral crest and the Pseudhesperosuchus but in this total length of humerus in Neuquensuchus crocodylomorph the posterior border is much wider universitas is 23.5%. In Sichuanosuchus it is also and it borns abruptly and more centrally (BONAPARTE, 23.5% and in Shantungosuchus it is 23% (WU et 1971). However, in Junggarsuchus (CLARK et al., 2004) al., 1997). This is different to the other the hook is dorsoposteriorly directed and the dorsal Metasuchia where this relationship is always border is slightly concave. bigger than 27%.

Fig.7- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-47. Sacral and first caudal vertebrae and left ilium. A and B, in left lateral view. C and E, sacral and left ilium in ventral view. D and F, sacral and left ilium in left lateral view. (Abbreviations in the Appendix IV).

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On the other , the diameter of the shaft in notosuchians. However, this characteristic is also relation to the total length of the humerus is, similar to Sichuanosuchus shuhanensis (WU et al., in Neuquensuchus (6.5%) similar to the other named 1997, IVPP V12088) and some Protosuchia and “protosuchians” (e.g., Sichuanosuchus, sphenosuchians, as for example Dibothrosuchus (WU Shantungosuchus, and Zaraasuchus), where it never & CHATTERJEE, 1993, IVPP V7907). This indicates that overcomes 7%, but this contrast with the the character in question does not throw mesoeucrocodylians Metasuchia where this overwhelming phylogenetic information because it relationship is always bigger than 9%. Moreover, in possesses a high distributional disparity inside Neuquensuchus universitas the relationship between Crocodylomorpha representing possible the total length and width of the proximal end of convergences in the different groups. However, the the humerus is approximately 5%, similar to those internal tuberosity of Neuquensuchus universitas is of Crocodylia, sphenosuchians, Protosuchia and more similar to that of Sichuanosuchus (WU et al., more basal crocodyliform, while in Metasuchia non- 1997). The lateral facet of the deltopectoral crest Crocodylia it is not bigger than 4%. has the border anterolaterally directed like in The proximal end of the humerus shows the articular Notosuchus and in the rest of the crocodyliforms it surface lateromedially elongated, strongly curved is laterally directed; however, in Sichuanosuchus medially and relatively thin anteroposteriorly, like shuhanensis (WU et al., 1997, IVPP V12088) this that present in Gobiosuchus and Sichuanosuchus lateral facet is seemingly also anterolaterally (Fig.10C). The lateroproximal expansion and the directed. rectangular proximal shape of the humerus of Distally, the medial condyle is bigger than the lateral Neuquensuchus universitas (Fig.10C) are very similar one and its general form and proportions are to those of Notosuchus (POL, 2005, MACN-RN 1037 identical to Sichuanosuchus shuhanensis (WU et al., and 1042), Chimaerasuchus paradoxus (WU & SUES, 1997). The posterolateral surface of the humerus is 1996, IVPP V8274), and patagonicus strongly concave and the posterior intercondylar (ORTEGA et al., 2000, MUCPv-267), suggesting some groove is broad, like in Sichuanosuchus huidongensis relationships between Neuquensuchus and these (PENG, 1996, ZDM 3404).

Fig.8- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-47. Left scapula in lateral view. (Abbreviations in the Appendix IV).

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Fig.9- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-47. Right humerus in anterior (A) and posterior (B) views. (Abbreviations in the Appendix IV).

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Fig.10- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-47. Left humerus in posterolateral (A), lateral (B) and anterior view (C). (Abbreviations in the Appendix IV).

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ULNA ILIUM

The ulna of Neuquensuchus universitas is straight Only the posterior fragment of the left ilium has been (Fig.11), with a long and thin shaft, slightly preserved in MUCPv-47 (Fig.7). It includes the compressed lateromedially as in Sichuanosuchus. posterior border of the acetabular cavity, the It possesses a small proximal expansion, and a ischiadic peduncle and postacetabular process. The convex surface for the lateral condyle of the posterior part of dorsal crest in Neuquensuchus humerus. As in other basal crocodylomorphs, like universitas is low and snub, different to Notosuchia some sphenosuchians but contrary to (POL, 2005; FIORELLI, 2005) where there is a very notosuchians, the ulna possesses a prominent laterally extended marked acetabular roof. The olecranon process. The right ulna, although length between the dorsal end of the crest and the incomplete, has a length of 107.5 mm and it is distal end of the ischiadic peduncle is very short, longer than the humerus. This character is only indicating an ilium dorsoventrally low. It differs from shared with some sphenosuchians (e.g., more derived Mesoeucrocodylia (Metasuchia) where Terrestrisuchus, Dibothrosuchus, and the ilium is very wide dorsoventrally. The ischiadic Dromicosuchus), representing an autapomorphy of peduncle is small and the surface for the articulation Neuquensuchus and a convergent feature shared of the ischium is reduced. The postacetabular with these sphenosuchians but related to the process is dorsoventrally thin and markedly cursorial habits of this crocodyliforms. However, posteriorly projected, with its distal extreme in Neuquensuchus universitas the relationship lateroventrally directed, like in Protosuchus (COLBERT between the width of the shaft (5.7mm) and their & MOOK, 1951) and other “protosuchian” forms. total length (107.5mm) is 5.3%; it is comparable to other “protosuchian” forms (Zaraasuchus <6%; PUBIS Gobiosuchus = 5%; Shantungosuchus = 5.6%; Sichuanosuchus = 5.3%) and differs from other The right pubis of Neuquensuchus universitas mesoeucrocodylians metasuchian where it is bigger (MUCPv-47) is practically complete. It is associated than 7% (notosuchians and neosuchians). to the proximal end of the right ischium, sacral and caudal vertebrae, left ilium and femur, tibia RADIUS and right fibula (Figs.12-13). The pubis is a long and thin bone (rod-like shaped), mainly in the The right radius (Fig.11) is a very long and thin bone. section of the shaft, similar to basal forms of It is similar in its general form to Sichuanosuchus Crocodylomorpha, as Terrestrisuchus (CRUSH, 1984), shuhanensis (WU et al., 1997, IVPP V12088). Its Protosuchus (COLBERT & MOOK, 1951), proximal end is strongly expanded and the thin shaft Sichuanosuchus (IVPP V12088), and a basal is circular in transverse section. The relationship innominated form of China (POL et al., 2004, between the diameter of the shaft (3.9mm) and total GMPKU-P 200102). The small proximal expansion length of the radius (105mm) in Neuquensuchus supports a convex facet for the ilium and for the universitas is 3.7%, which is similar to pubic process of the ischium (Fig.12B). This Sichuanosuchus shuhanensis (3.6%). By contrast in character is similar to that of Sichuanosuchus and Terrestrisuchus it is 2.9% and in the other other “protosuchians”, and it implies that the pubis sphenosuchians it is bigger (for example in is partially introduced inside the acetabulum. The Pseudhesperosuchus it is 5% and in pubis is slightly expanded distally, as in GMPKU- it is 5.75%). In more derived members of P 200102 (POL et al., 2004) and Sichuanosuchus. Mesoeucrocodylia this relationship always In Neuquensuchus universitas the relationship surpasses 5% (Araripesuchus patagonicus: 5.5%; between the length of the pubis (39.5mm) and the Notosuchus: 8.05%; Chimaerasuchus: >8%; width of the distal expansion (10.8mm) is 27%, : >8%; Crocodylia: = 8%) contrary to similar to Sichuanosuchus (26%) and Gobiosuchus Araripesuchus tsangatsangana (TURNER, 2006) where (23-24%). In more derived Mesoeucrocodylia – it is 4.52%. metasuchian forms –, this proportion is always The specimen MUCPv-47 possesses a small superior to 30%. Lastly, the diameter of the pubic proximal fragment of the radial, articulated to the shaft, in relation to the total length, resembles that end of the right radius, which is very similar to of other “protosuchians”. In Neuquensuchus, this Sichuanosuchus shuhanensis (WU et al., 1997). relationship is 7%, similar to Sichuanosuchus

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(6.5%) and Gobiosuchus (<7%) and very different been preserved in MUCPv-47, together with a from Metasuchia (>8%). slight impression (Fig.12B). It is very similar in The existent relationship between the total length its construction to Protosuchus, Sichuanosuchus of the pubis and the total length of femur is a and GMPKU-P 200102 (POL et al., 2004). The characteristic only shown by Gobiosuchus, pubis process of ischium is slightly narrower than Shantungosuchus and Neuquensuchus being smaller the proximal end of the pubis, like in than 45%, while in Terrestrisuchus, Protosuchus, and Sichuanosuchus, and it contacts with the Metasuchia the proportion between pubis and femur pubis in its posterodorsal extreme. For this is always bigger due mainly to the reduction of the reason, the ischium partially excludes the pubis, to exception of Mahajangasuchus. pubis of the acetabulum. The half section of the proximal shaft shows that it is quite narrow

ISCHIUM but it spreads to distally expanded according to the impression of the same similar to Protosuchus Only the proximal end of the right ischium has and Gobiosuchus.

Fig.11- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-47. Right ulna, radius and radial in lateral (A and C) and medial (B and D) views. (Abbreviations in the Appendix IV).

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Fig.12- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-47. A, right pubis and femur in lateral view; B, right pubis, ischium and femur in medial view. (Abbreviations in the Appendix IV).

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FEMUR the femoral fragment of Shantungosuchus hangjinensis (WU et al., 1994, IVPP V10097). Only in the specimen MUCPv-47 of Neuquensuchus Neuquensuchus universitas as in other basal universitas the right femur have been preserved crocodyliforms lacks of a prominent anteromedial (Figs.3D, 13, 14). In the holotype, the right femur process of the femur medially placed on the articulates with the tibia and fibula (Figs.12-13) proximal end of shaft. This process is very marked as likewise with the right ilum, sacral and first in Notosuchia (POL, 2005; FIORELLI, 2005; fig. 14B) caudals vertebrae. The long and thin femur is and other metasuchians such as like in basal crocodylomorphs. It is mostly Mahajangasuchus (BUCKLEY & BROCHU, 1999). practically straigth and the sigmoid form is not Although in MUCPv-47 the distal end is damaged conspicuous or not very marked. The condyle on we can observe that the lateral condyle (fibular the femoral head is slightly expanded (Fig.14). c.) is slightly bigger with respect to the medial This characteristic differs from other one. An important character in Neuquensuchus sphenosuchians, such as Terrestrisuchus (CRUSH, is the relationship of the diaphyseal width (7mm) 1984), Dromicosuchus (SUES et al., 2003, UNC and the total length of the femur (94mm) equal to 15574), Macelognathus (GÖHLICH et al., 2005, LACM 7.5%. This is similar to some basal crocodyliforms 4684/128272), and derived mesoeucrocodylians. (Gobiosuchus = 6.3%; Shantungosuchus = 7.6%), The femur of Neuquensuchus universitas possesses differing from Protosuchus and more derived a lengthened furrow similar in its proportions mesoeucrocodylians – Metasuchia – where it is and muscular dispositions to that observed in always bigger than 9%.

Fig.13- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-47. Right pubis, femur, tibia and fibula in lateral view. (Abbreviations in the Appendix IV).

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TIBIA length. This possibly represents one of the most important characters in the species because this The right tibia in MUCPv-47 is complete (Fig.13), while feature character is only shared with in MUCPv-161 just the proximal end is preserved Shantugosuchus, where the length of the femur is (Fig.14). The tibia possesses a very long, straigth and 95% of the tibial length (WU et al., 1994) (see Fig.13). thin shaft, similar to that present in some most basal By contrast in all other crocodyliforms the femur Crocodylomorpha, as in sphenosuchians like is always longer than the tibia (WU et al., 1994). Macelognathus, Dromicosuchus, Hesperosuchus, and Even so, in early ontogenetic states of Crocodylia Terrestrisuchus (CRUSH, 1984; CLARK et al., 2000; SUES the femur is always longer than the tibia et al., 2003; GÖHLICH et al., 2005). However, in some (DODSON, 1975). Inside Crocodylomorpha, some “protosuchian” forms the tibia is too similar, such as sphenosuchians as Terrestrisuchus or in Shantungosuchus chuhsienensis (YOUNG, 1961; WU Macelognathus have the tibia longer than the femur et al., 1994, IVPP V2484) and Gobiosuchus kielanae (SERENO, 1991; CRUSH, 1984; GÖHLICH et al., 2005). (OSMÓLSKA, 1972; OSMÓLSKA et al., 1997, ZPAL MgR- On the other hand, the relationship between the II/67). The proximal end is broad and the distal end diaphyseal width (5.6mm) and the tibia length has a small lateromedial expansion. Neuquensuchus (105mm; 5.3%) is identical to that of universitas does not possess a developed cnemial crest Shantungosuchus (5.3%), differing from those of and the femoral condyles form a marked notch in Protosuchus and Metasuchia that is always bigger the distal end (Figs.14A, 14C). then 8%. The discussions and evolutionary In MUCPv-47, the tibia (105.3mm) is longer than consequences on these characteristics are offered the femur (94.5mm) comprising 89.7% of the tibial later on (see Discussion).

Fig.14- Neuquensuchus universitas gen.nov., sp.nov., MUCPv-161. A, B and C, proximal end of left tibia; D and E, distal end of left fibula. Tibia in posterior (A), anterior (B) and lateral (C) view. Fibula in medial (D) and lateral (E) views. (Abbreviations in the Appendix IV).

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FIBULA neosuchian group. However, Notosuchia as well as the basal groups of crocodyliformes stayed Few fibular materials have been preserved. In constant in the different hypotheses as we can MUCPv-47, the partial and very fragmentary right observe in the strict consensus tree (Fig.15). fibula (Fig.13) is thin and long. In MUCPv-161 the In all more parsimonious hypotheses, distal end of fibula possesses a thin shaft with D- Neuquensuchus universitas represents the sister shaped in cross-section (Figs.14D, 14E). taxa of Shantungosuchus hangjinensis from the Lower Cretaceous of Inner Mongolia (Northern TARSUS China). Both shared the character 91 “hypapophyses present only in cervical vertebrae” Only conserved in MUCPv-161, the left astragalus and character 226 “Tibia longer than the femur” is incompletely preserved (Fig. 3D). In spite of it, (Node 11 of the figure 15). This last character is we can see morphological characters in the ambiguous in Sichuanosuchus shuhanensis and articulations that are present in typical Zosuchus davidsoni. In another sence, just two Crocodyliformes tarsus. For instance, a good diagnostic character separating Neuquensuchus marked process supporting the square fibular facet from Shantungosuchus (olecranon well developed and a lateromedially wide tibial facet. The articulate [character 173-0] and the relationship between the surface for the metatarsals is rounded and width ulna length and the humerus length [Character with a deep anterior hollow. 220]; Node 12). The absence of additional autapomophies in Neuquensuchus can be due to the fragmentarity of the available material, which PHYLOGENETIC RELATIONSHIPS does not possess cranial remains, the reason why we support the erection of this new taxon. The Although Neuquensuchus universitas gen.nov., temporal and geographical separation goes in favor sp.nov. is represented just by postcranials of this proposal. The resulting shows that remains, it was possible to establish its Neuquensuchus and Shantungosuchus are the phylogenetic relationships based on parsimony of Sichuanosuchus shuhanensis from analysis. For this analysis, we used a modified the of Sichuan, China (Node 10). data set taken of recent publications (POL & NORELL, This node is diagnosed by two unambiguous 2004b; POL et al., 2004), which was based on the synapomorphies (palatines form palatal shelves addition of several characters of previously that do not meet [Character 37]; posteroventral edge published matrix (CLARK, 1994; WU & SUES, 1996; of mandibular ramus markedly deflected GOMANI, 1997; WU et al., 1997; BUCKLEY et al., 2000; [Character 170]). However, both characters are ORTEGA et al., 2000). We have included new ambiguous in Neuquensuchus. Zosuchus davidsoni characters not included in previous publications from Upper Cretaceous of Gobi Desert (Mongolia), that were defined by WU & SUES (1996), MARTINELLI represents the sister taxa of the resulting node of (2003), and FIORELLI (2005). Moreover, sixteen new the three previously taxa (Node 9), diagnosed by characters were added and new taxa were five unambiguous synapomorphies (characters 55, included. The matrix includes 231 characters and 143, 163, 169 and 178; see Appendix I). 51 taxa (see appendixes I and II). The present work The clade conformed by Fruita form, Zosuchus, tries to focus mainly in non-neosuchian basal Sichuanosuchus, Shantungosuchus, and crocodyliforms. In the present analysis, characters Neuquensuchus (Node 8 from the figure 15), is were taken with equal weight using NONA closely related to Hsisosuchus and more derived (GOLOBOFF, 1993) and published with Winclada mesoeucrocodilians than other Protosuchia (NIXON, 1999). An heuristic tree search was (Gobiosuchus, Protosuchus and all their performed consisting of 1000 replicates of RAS + descendants). This conclusion is similar to that TBR with a final round of TBR (mult*1000; max*;), obtained in other works (POL, 2003; POL & NORELL, holding 20 trees per replication (hold/20;). Thirty 2004a, 2004b; POL et al., 2004; FIORELLI, 2005; POL six (36) most parsimonious trees of 839 steps (CI & APESTEGUÍA, 2005; ZAHER et al., 2006), but differs 0.34; RI 0.65) were found in all of replications. of those in that it postulates a monophyly of The 36 phylogenetic hypotheses differ in the protosuchids and “protosuchians” (e.g., WU et al., relationships of some neosuchian crocodyliforms 1994; WU & SUES, 1996; WU et al., 1997; TYKOSKI et like for instance Peirosaurid forms and derived al., 2002).

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Fig.15- Strict consensus of the 36 most parsimonious topologies that resulted from a strict parsimony analysis obtained with NONA and published with Winclada. Tree length is 839 with a CI of .33 and a RI of .65. The tree shows the phylogenetic relationships of Neuquensuchus universitas gen.nov., sp.nov. performed a basal mesoeucrocodylia. 1: Crocodylomorpha; 2: “Sphenosuchia”; 3: Crocodyliformes; 4: Protosuchia; 5: Gobiosuchidae; 6: ; 7: Mesoeucrocodylia; 8, 9, 10 and 11: Innominated; 12: Neuquensuchus universitas gen.nov., sp.nov.; 13: “”; 14: Metasuchia; 15: ; 16: ; 17: ; 18 and 19: Innominated; 20: Notosuchia; 21: ; 22: Innominated; 23: Notosuchidae; 24: . Araripesuchus is used here like a terminal taxon although in the analyses it was used A. gomesii and A. patagonicus. Explanation and definitions of suprageneric taxa see Appendix V.

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On the other hand and in contrast to some recent example Lissamphibia (Discoglossidae, phylogenetic analysis (e.g., CLARK, 1994; BUCKLEY Callobatrachus), Mammaliamorpha (e.g., et al., 2000; ORTEGA et al., 2000; TURNER, 2004; Peramura), Notosuchia (Chimaerasuchus) and 2006; TURNER & C ALVO, 2005) that placed (cf. Theriosuchus). Even in countless Araripesuchus like basal member of Neosuchia, groups of dinosaurs, for example in our analyses this taxon appears as a basal Rebbachisauridae, Nemegtosauridae, member of notosuchian clade, an important result Saltasauridae, Abelisauroidea, Spinosauroidea, comparable with other recent phylogenetic studies Carcharodontosauridae, Deinonychosauria, (POL, 2003; POL & NORELL, 2004a, 2004b; POL et Alvarezsauria, and some Ornithischia al., 2004; FIORELLI, 2005; POL & APESTEGUÍA, 2005; (Valdosaurus, ). Summing up, it was ZAHER et al., 2006). suggested by different authors (e.g., WU & SUES, 1996; POL, 2003), that this rises many questions to the hypothesis of faunistic endemism in DISCUSSION Gondwana during Cretaceous times, a classic hypothesis assumed by several authors (GASPARINI, Neuquensuchus universitas gen.nov., sp.nov. 1971; BONAPARTE, 1986; 1991; CLARK et al., 1989). represents the first basal Mesoeucrocodylia non The occurrence of Neuquensuchus in Gondwana Metasuchia from the Cretaceous, not only from does not indicate the presence of Pangeic lineage Argentina but also from South America and of this clade in Southern lands. The presence of Gondwana (Fig.16). Mesoeucrocodylia is defined this basal mesoeucrocodylian is more probably due here like the most inclusive clade containing to subsequent dispersion, as it has been postulated but not Protosuchus (BENTON & CLARK, in recent studies by JUÁREZ VALLIERI & FIORELLI (2002; 1988; CLARK, 1994; sensu SERENO et al., 2001; 2005). 2003) and FIORELLI (2005). These authors propose Without doubts, the Triassic argentinean and a dispersion event among Gondwana, Europe and gondwanic forms of basal crocodyliforms, such as Central Asia during the Early Cretaceous Hemiprotosuchus (B ONAPARTE, 1967; 1971), (–), producing a faunistic Protosuchus sp. (ALCOBER et al., 2004), Orthosuchus interchange poorly recognized previously (BRETT- (NASH, 1975), and Baroqueosuchus haughtoni SURMAN, 1979). Probably it occurred in both ways: (BUSBEY & GOW, 1984) are not related directly to from Central Asia to Gondwana through Europe Neuquensuchus universitas, because it integrates as well as in the opposite direction. This new the most basal group of Mesoeucrocodylia (Figs.15- hypothesis agrees with the distributional pattern 16) due to the intimate relationships with other so of all groups and is perfectly adjusted with formerly called “protosuchians” and more derived recent genetic studies carried out on current form like Hsisosuchus. Then, it demonstrates that vertebrates (see HAY et al., 1995; HEDGES & POLING, Neuquensuchus does not represent a derived form 1999; HEDGES, 2001; COOPER et al., 2001; MURPHY et from the Upper Triassic/Early Gondwana al., 2001; MEYER & ZARDOYA, 2003). taxa. Therefore, it comes from highly more derived These basals mesoeucrocodylian non-Metasuchia taxa from the Early Cretaceous of Central Asia, were abundant during Jurassic and Cretaceous in such as Shantungosuchus and Sichuanosuchus Asia. Undoubtedly they came from basal forms of (Figs. 15, 16). It would be possible that related form Upper Triassic or times, which of “Las Hoyas crocodyliform” is closely related, but have suffered an adaptative radiation in that there is not a detailed data of this specimen to continent. Posteriorly in the Early Cretaceous, include it in the phylogenetic analyses, although after the contact between Gondwana and Asia in recent studies “Las Hoyas crocodyliform” is (JUÁREZ VALLIERI & FIORELLI, 2002; 2003; FIORELLI, intimately related to Gobiosuchus (see ORTEGA et al., 2005), dispersion toward Southern continents 2000). Regarding to the Cretaceous of well derived forms took place and for this paleobiogeography, Neuquensuchus universitas reason Neuquensuchus universitas occurs in throws more problems than answers inside the Northern Patagonia. Summing up, Neuquensuchus classics paleogeographic models used until now represents a clade of mesoeucrocodylian basal (e.g., BONAPARTE, 1986; BUFFETAUT, 1982; SERENO, form with a purely Asian origin and dispersal 1999). This problematic disjunct distributional center, at least during the Upper Jurassic, Cretaceous pattern is similar to that observed in and with dispersion out of Asia toward Europe other groups of very diverse tetrapods, as for and Gondwana during the Early Cretaceous.

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Fig.16- Chronological distribution of Crocodylomorpha. The “Shartegosuchidae” (EFIMOV, 1988) and other taxa not included in the phylogenetics analysis alone indicating here the highly endemic fauna of Crocodyliformes present in Central Asia during Jurassic and Cretaceous times; they do not indicate phylogenetic relationships with other groups in this chronology.

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In the opposite way, we can explain the presence like Terrestrisuchus or Macelognathus the tibia is in the Early Cretaceous of China of the derived longer than the femur (SERENO, 1991; CRUSH, 1984; notosuchian Chimaerasuchus paradoxus (WU & GÖHLICH et al., 2005). Undoubtedly, this convergent SUES, 1996; MARTINELLI, 2003; POL, 2003; POL & characteristic was acquired independently by both NORELL, 2004a; 2004b; POL, 2005; FIORELLI, 2005) groups, sphenosuchians – some species – and these or the atoposarid neosuchian cf. Theriosuchus sp. two basal mesoeucrocodylian taxa, (WU et al., 1996, IVPP V 10613). It seems that Shantungosuchus and Neuquensuchus. derived basal Crocodyliformes had an important As it has been suggested by diverse authors (CRUSH, adaptative success in Central Asia during 1984; SERENO, 1991; SERENO & WILD, 1992; CLARK et Cretaceous times, for example by the occurrence al., 2000; SUES et al., 2003; CLARK et al., 2004; GÖHLICH of Edentosuchus, Tagarosuchus, Artzosuchus, et al., 2005), sphenosuchians such as Gobiosuchus, Zaraasuchus, Shantungosuchus, Terrestrisuchus, Macelognathus, Junggarsuchus, Sichuanosuchus, and Zosuchus. By contrast, in Dromicosuchus, and Hesperosuchus, would have Neopangea it did not happen this way. The fact presented a high capacity cursorial for the diverse that in Gondwana, and mainly in South America, characteristics of their extremities, mainly by the exist an acceptable Cretaceous crocodyliform record, long and thin bones. Also, WU et al. (1994) suggested the fragmentary remains of Neuquensuchus probably that Shantungosuchus had a high cursorial capacity indicate their low abundance. Moreover they did not instead of very quick terrestrial displacement. The suffer an apparent adaptative radiation, as it close relationships of forelimb with Neuquensuchus occurred with Notosuchia, a properly gondwanic allow us to expect the same capacity of movement group. Together with previous protosuchid and and cursorial capacity. In the more related taxa “protosuchians” Asian taxa, the Upper Jurassic and (Sichuanosuchus and Zosuchus), this characteristic Early Cretaceous forms includes in – tibia > femur – is ambiguous. These important “Shartegosuchidae” – Shartegosuchus (EFIMOV, 1988), cursorial characteristic present in these Nominosuchus (EFIMOV, 1996; KURZANOV et al., 2003), crocodylomorphs possibly had a great influence in Kyasuchus (EFIMOV & LESHCHINSKIY, 2000), and their spatial ranges and the amplification of Adzosuchus (EFIMOV et al., 2000) (see Fig.16) –, it is ecological and territorial niches, allowing a bigger indicating the highly endemic fauna of dispersal capacity. Crocodyliformes present in Central Asia during Although postcranials remains of Neuquensuchus Jurassic and Cretaceous times. In a recent work universitas gen.nov., sp.nov. reported here represent (FIORELLI et al., 2006), it was demonstrated the the first evident crocodyliform non-Metasuchia in monophyly and the Shartegosuchidae’s endemic gondwanic Cretaceous lands, we do not know too group, and they represent the most basal group of much about their anatomy and relationships. We the mesosuchian clade (Fiorelli et al., in prep.). The believe that the strong phylogenetic relationships crocodyliforms fauna and other Asian continental of Neuquensuchus produce important implications tetrapods are correlated with the biogeographical and give novel light about the paleobiogeographic hypothesis proposed by RUSSELL (1993) of a issues. New exploratory works with the purpose of sequential partition of Pangea. He postulated the finding new remains of these original taxa, mainly Asian isolation of Neopangea during the Upper cranial materials, will help to elucidate and know Triassic or Early Jurassic. with more details their anatomy and phylogenetic In another sense, an interesting aspect that relationships. presents Neuquensuchus universitas derivated from the present study is the important character related to the longitudinal ratio between the femur and ACKNOWLEDGEMENTS tibia. Previously it was aforementioned that the femur comprises 89.7% of the tibial length, feature We want to express our most sincere gratefulness character only shared with Shantugosuchus, with to the technicians of the Lake Barreales a femoral length of 95% of the total tibial length Paleontological Center for partly preparation of the (WU et al., 1994) (see Fig.13). In the other material; to the members of the CePaLB for the crocodyliforms the femur is always longer than the colaboration during the study of the material. tibia (WU et al., 1994), even so in early ontogenetics Additional gratefulness to MSc. Marco Brandalise states of Crocodylia (DODSON, 1975). Within de Andrade and to Dr. Diego Pol is here expressed Crocodylomorpha, only in some sphenosuchians for the informations given and the valuable

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WOODWARD, A.S., 1896. On two Mesozoic crocodilians WU, X.-C.; SUES, H.-D. & BRINKMAN, D.B., 1996. An Notosuchus ( novum) and (genus atoposaurid neosuchian (Archosauria: Crocodyliformes) novum), from the Red Sandstones of the Territory of from the Lower Cretaceous of Inner Mongolia (People’s Neuquén (Argentina Republic). Anales del Museo de La Republic of China). Canadian Journal of Earth Plata, Paleontología, 4:1-20. Sciences, 33:599-605.

WOODWARD, A.S., 1901. On some extinct reptiles from WU, X.-C; SUES, H.-D. & DONG, Z.-M., 1997. Patagonia of the genera Miolana, Dinilysia and Sichuanosuchus shuhanensis, a new ?Early Cretaceous Genyodectes. Proceedings of the Zoological Society protosuchian (Archosauria: Crocodyliformes) from of London, 1:169-184. Sichuan (China), and the monophyly of Protosuchia. Journal of Vertebrate Paleontology, 17:89-103. WU, X.-C.; BRINKMAN, D.B. & LU, J.-C., 1994. A new species of Shantungosuchus from the Lower Cretaceous YOUNG, C.C., 1961. Shantungosuchus chuhsienensis, of Inner Mongolia (China), with comments on S. a new crocodile. Vertebrata PalAsiatica, 5:6-15. chuhsienensis Young, 1961, and the phylogenetic position of the genus. Journal of Vertebrate YOUNG, C.C., 1973. A new fossil crocodile from Wuerho. Paleontology, 14:210-229. Memoirs Institute Vertebrate Paleontology and Paleoanthropology, 11:37-44. WU, X.-C. & CHATTERJEE, S., 1993. Dibothrosuchus elaphros, a crocodylomorph from the Lower Jurassic of YOUNG, C.C. & CHOW, M.C., 1953. New discovery of China and the phylogeny of the Sphenosuchia. Journal Reptilia from the Mesozoic of Sichuan. Acta of Vertebrate Paleontology, 13:58-89. Palaeontologica Sinica, New Series C, 1:87-111.

WU, X.-C. & SUES, H.-D., 1996. Anatomy and ZAHER, H.; POL, D; CARVALHO, A.B.; RICOMINI, C.; phylogenetic relationships of Chimaerasuchus CAMPOS, D. & NAVA, W., 2006. Redescription of the paradoxus, an unusual crocodyliform reptile from the cranial morphology of Mariliasuchus amarali, and its Lower Cretaceous of Hubei, China. Journal of phylogenetic affinities (Crocodyliformes, Notosuchia). Vertebrate Paleontology, 16:688-702. American Museum Novitates, 3512:1-40.

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APPENDIX I

LISTS OF CHARACTERS CORRESPONDING TO THE DATA MATRIX (SEE APPENDIX III) USED IN THE PHYLOGENETIC ANALYSES

Definitions of the characters 1-101 were taken from CLARK (1994) and they have the same numeration like in the original publication. The character 5 was excluded of this analysis (due to the dependence with the modified definition of the character 6). Nevertheless, this exclusion does not affect the result of these analyses. The following ones, 102 to 192 characters, were taken from POL & NORELL (2004b). They are listed in order in relation to the same publication and the source mentioned together with the number of original character. The characters 193 and 194 were taken and designated by POL et al. (2004), corresponding originally to the characters 164 and 179, respectively. The characters 195, 196, and 197 were taken from WU & SUES (1996) that originally corresponded to the characters 6, 17 and 31, respectively. Although the characters 198 and 199 were taken from MARTINELLI (2003) they originally corresponded to the respective characters 35 and 36. The characters 200 and 210 were designated by FIORELLI (2005) and the numerations are the same ones. The characters 215 and 218 were taken and modified from POL (1999a) corresponding to the characters 192 and 191, respectively. Character 226 is taken and modified from SERENO (1991) corresponding to the character 27. The characters 1, 3, 6, 23, 37, 45, 49, 65, 67, 69, 73, 77, 79, 90, 91, 96, 97, 103, 104, 105, 107, 126, 143, 149, and 165 were taken as aditives characters (also marked with “+” in this list). For finish, the characters 211– 214, 216, 217, 219–225, and 227–231 are new, designated by the authors.

CHARACTER 1 (modified from CLARK, 1994: character 1): + External surface of dorsal cranial bones: smooth (0), slightly grooved (1) and heavily ornamented with deep pits and grooves (2). CHARACTER 2 (modified from CLARK, 1994: character 2): Skull expansion at orbits: gradual (0), or abrupt (1). CHARACTER 3 (modified from CLARK, 1994: character 3): + Rostrum proportions: narrow oreinirostral (0), broad oreinirostral (1), nearly tubular (2), or platyrostral (3). CHARACTER 4 (CLARK, 1994: character 4): Premaxilla participation in internarial bar: forming at least the ventral half (0), or with little participation (1). CHARACTER 5 (CLARK, 1994: character 5): Premaxilla anterior to nares: narrow (0), or broad (1). CHARACTER 6 (modified from CLARK, 1994: character 6): + External nares facing anterolaterally or anteriorly (0), dorsally not separated by premaxillary bar from anterior edge of rostrum (1), or dorsally separated by premaxillary bar (2). CHARACTER 7 (CLARK, 1994: character 7): Palatal parts of premaxillae: do not meet posterior to incisive foramen (0), or meet posteriorly along contact with maxillae (1). CHARACTER 8 (CLARK, 1994: character 8): Premaxilla- contact: premaxilla loosely overlies maxilla (0), or sutured together along a butt joint (1). CHARACTER 9 (modified from CLARK, 1994: character 9): Ventrally opened notch on ventral edge of rostrum at premaxilla- maxilla contact: absent (0), present as a notch (1), or present as a large fenestra (2). CHARACTER 10 (CLARK, 1994: character 10): Posterior ends of palatal branches of maxillae anterior to palatines: do not meet (0), or meet (1). CHARACTER 11 (CLARK, 1994: character 11): Nasal contacts lacrimal (0), or does not contact (1). CHARACTER 12 (CLARK, 1994: character 12): Lacrimal contacts nasal along medial edge only (0), or medial and anterior edges (1). CHARACTER 13 (CLARK, 1994: character 13): Nasal contribution to narial border: yes (0), or no (1). CHARACTER 14 (CLARK, 1994: character 14): Nasal-premaxilla contact: present (0), or absent (1). CHARACTER 15 (modified from CLARK, 1994: character 15): Descending process of prefrontal: does not contact palate (0), or contacts palate (1). CHARACTER 16 (CLARK, 1994: character 16): Postorbital-jugal contact: postorbital anterior to jugal (0), or postorbital medial to jugal (1), or postorbital lateral to jugal (2). CHARACTER 17 (CLARK, 1994: character 17): Anterior part of the jugal with respect to posterior part: as broad (0), or twice as broad (1). CHARACTER 18 (CLARK, 1994: character 18): Jugal bar beneath infratemporal fenestra: flattened (0), or rod-shaped (1). CHARACTER 19 (CLARK, 1994: character 19): Quadratojugal dorsal process: narrow, contacting only a small part of postorbital (0), or broad, extensively contacting the postorbital (1). CHARACTER 20 (CLARK, 1994: character 20): Frontal width between orbits: narrow, as broad as nasals (0), or broad, twice as broad as nasals (1). CHARACTER 21 (CLARK, 1994: character 21): Frontals: paired (0), unpaired (1). CHARACTER 22 (CLARK, 1994: character 22): Dorsal surface of frontal and parietal: flat (0), or with midline ridge (1). CHARACTER 23 (modified from CLARK, 1994: character 23 by BUCKLEY & BROCHU, 1999: character 81): + Parieto-postorbital suture: absent from dorsal surface of skull roof and supratemporal fossa (0), absent from dorsal surface of skull roof but broadly present within supratemporal fossa (1), or present within supratemporal fossa and on dorsal surface of skull roof (2).

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CHARACTER 24 (CLARK, 1994: character 24): Supratemporal roof dorsal surface: complex (0), or dorsally flat “skull table” developed, with postorbital and squamosal with flat shelves extending laterally beyond quadrate contact (1). CHARACTER 25 (modified from CLARK, 1994: character 25) Postorbital bar: sculpted (if skull sculpted) (0), or unsculpted (1). CHARACTER 26 (modified from CLARK, 1994: character 26): Postorbital bar: transversely flattened (0), or cylindrical (1). CHARACTER 27 (CLARK, 1994: character 27): Vascular opening in dorsal surface of postorbital bar: absent (0), or present (1). CHARACTER 28 (modified from CLARK, 1994: character 28): Postorbital anterolateral process: absent or poorly developed (0), or well developed, long, and acute (1). CHARACTER 29 (CLARK, 1994: character 29): Dorsal part of the postorbital: with anterior and lateral edges only (0), or with anterolaterally facing edge (1). CHARACTER 30 (CLARK, 1994: character 30): Dorsal end of the postorbital bar broadens dorsally, continuous with dorsal part of postorbital (0), or dorsal part of the postorbital bar constricted, distinct from the dorsal part of the postorbital (1). CHARACTER 31 (CLARK, 1994: character 31): Bar between and supratemporal fossa broad and solid, with broadly sculpted dorsal surface (0), or bar narrow, sculpting restriced to anterior surface (1). CHARACTER 32 (modified from CLARK, 1994: character 32): Parietal: with broad occipital portion (0), or without broad occipital portion (1). CHARACTER 33 (CLARK, 1994: character 33): Parietal: with broad sculpted region separating fossae (0), or with sagittal crest between supratemporal fossae (1). CHARACTER 34 (CLARK, 1994: character 34): Postparietal (dermosupraoccipital): a distinct element (0), or not distinct (fused with parietal?) (1). CHARACTER 35 (CLARK, 1994: character 35): Posterodorsal corner of the squamosal: squared off, lacking extra “lobe” (0), or with unsculptured “lobe” (1). CHARACTER 36 (modified from CLARK, 1994: character 36): Posterolateral process of squamosal: poorly developed and projected horizontally at the same level of the skull (0), elongated, thin, and posteriorly directed, not ventrally deflected (1), or elongated, posterolaterally directed, and ventrally deflected (2). CHARACTER 37 (CLARK, 1994: character 37): + Palatines: do not meet on palate below the narial passage (0), form palatal shelves that do not meet (1), or meet ventrally to the narial passage, forming part of secondary palate (2). CHARACTER 38 (CLARK, 1994: character 38): Pterygoid: restricted to palate and suspensorium, joints with quadrate and basisphenoid overlapping (0), or pterygoid extends dorsally to contact laterosphenoid and form ventrolateral edge of the trigeminal foramen, strongly sutured to quadrate and laterosphenoid (1). CHARACTER 39 (modified from CLARK, 1994: character 39): Choanal opening: continuous with pterygoid ventral surface except for anterior and anterolateral borders (0), or opens into palate through a deep midline depression (choanal groove) (1). CHARACTER 40 (CLARK, 1994: character 40): Palatal surface of pterygoids: smooth (0), or sculpted (1). CHARACTER 41 (CLARK, 1994: character 41): Pterygoids posterior to choanae: separated (0), or fused (1). CHARACTER 42 (modified from CLARK, 1994: character 42 by ORTEGA et al., 2000: character 139): Depression on primary pterygoidean palate posterior to choana: absent or moderate in size being narrower than palatine bar (0), or wider than palatine bar (1). CHARACTER 43 (CLARK, 1994: character 43): Pterygoids: do not enclose choana (0), or enclose choana (1). CHARACTER 44 (modified from CLARK, 1994: character 44): Anterior edge of choanae situated near posterior edge of suborbital fenestra (or anteriorly) (0), or near posterior edge of pterygoid flanges (1). CHARACTER 45 (CLARK, 1994: character 45): + Quadrate: without fenestrae (0), with single fenestrae (1), or with three or more fenestrae on dorsal and posteromedial surfaces (2). CHARACTER 46 (CLARK, 1994: character 46): Posterior edge of quadrate: broad medial to tympanum, gently concave (0), or posterior edge of quadrate narrow dorsal to otoccipital contact, strongly concave (1). CHARACTER 47 (CLARK, 1994: character 47): Dorsal, primary head of quadrate articulates with squamosal, otoccipital, and prootic (0), or with prootic and laterosphenoid (1). CHARACTER 48 (CLARK, 1994: character 48): Ventrolateral contact of otoccipital with quadrate: very narrow (0), or broad (1). CHARACTER 49 (CLARK, 1994: character 49): + Quadrate, squamosal, and otoccipital: do not meet to enclose cranioquadrate passage (0), enclose passage near lateral edge of skull (1), or meet broadly lateral to the cranioquadrate passage (2). CHARACTER 50 (CLARK, 1994: character 50): Pterygoid ramus of quadrate: with flat ventral edge (0), or with deep groove along ventral edge (1). CHARACTER 51 (CLARK, 1994: character 51): Ventromedial part of quadrate: does not contact otoccipital (0), or contacts otoccipital to enclose carotid artery and form passage for cranial nerves IX–XI (1). CHARACTER 52 (CLARK, 1994: character 52): Eustachian tubes: not enclosed between basioccipital and basisphenoid (0), or entirely enclosed (1). CHARACTER 53 (CLARK, 1994: character 53): Basisphenoid rostrum (cultriform process): slender (0), or dorsoventrally expanded (1).

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CHARACTER 54 (CLARK, 1994: character 54): Basipterygoid process: prominent, forming movable joint with pterygoid (0), or basipterygoid process small or absent, with basisphenoid joint suturally closed (1). CHARACTER 55 (modified from CLARK, 1994: character 55 by ORTEGA et al., 2000: character 68): Basisphenoid ventral surface: shorter than the basioccipital (0), or wide and similar to, or longer in length than basioccipital (1). CHARACTER 56 (CLARK, 1994: character 56): Basisphenoid: exposed on ventral surface of braincase (0), or virtually excluded from ventral surface by pterygoid and basioccipital (1). CHARACTER 57 (CLARK, 1994: character 57): Basioccipital: without well-developed bilateral tuberosities (0), or with large pendulous tubera (1). CHARACTER 58 (CLARK, 1994: character 58): Otoccipital: without laterally concave descending flange ventral to subcapsular process (0), or with flange (1). CHARACTER 59 (CLARK, 1994: character 59): Cranial nerves IX–XI: pass through common large foramen vagi in otoccipital (0), or cranial nerve IX passes medial to nerves X and XI in separate passage (1). CHARACTER 60 (CLARK, 1994: character 60): Otoccipital: without large ventrolateral part ventral to paroccipital process (0), or with large ventrolateral part (1). CHARACTER 61 (CLARK, 1994: character 61): Crista interfenestralis between fenestrae pseudorotunda and ovalis nearly vertical (0), or horizontal (1). CHARACTER 62 (CLARK, 1994: character 62): Supraoccipital: forms dorsal edge of the foramen magnum (0), or otoccipitals broadly meet dorsal to the foramen magnum, separating supraoccipital from foramen (1). CHARACTER 63 (CLARK, 1994: character 63): Mastoid antrum: does not extend into supraoccipital (0), or extends through transverse canal in supraoccipital to connect middle ear regions (1). CHARACTER 64 (CLARK, 1994: character 64): Posterior surface of supraoccipital: nearly flat (0), or with bilateral posterior prominences (1). CHARACTER 65 (modified from CLARK, 1994: character 65): + One small palpebral present in orbit (0), one large palpebral (1), or two large palpebrals (2). CHARACTER 66 (CLARK, 1994: character 66): External nares: divided by a septum (0), or confluent (1). CHARACTER 67 (CLARK, 1994: character 67): + Antorbital fenestra: as large as orbit (0), about half the diameter of the orbit (1), much smaller than the orbit (2), or absent (3). CHARACTER 68 (modified from CLARK, 1994: character 68 by ORTEGA et al., 2000: character 41): Supratemporal fenestrae extension: relatively large, covering most of surface of skull roof (0), or relatively short, fenestrae surrounded by a flat and extended skull roof (1). CHARACTER 69 (modified from CLARK, 1994: character 69): + Choanal groove: undivided (0), partially septated (1), or completely septated (2). CHARACTER 70 (CLARK, 1994: character 70): Dentary: extends posteriorly beneath mandibular fenestra (0), or does not extend beneath mandibular fenestra (1). CHARACTER 71 (modified from CLARK, 1994: character 71): Retroarticular process: absent or extremely reduced (0), very short, broad, and robust (1), with an extensive rounded, wide, and flat (or slightly concave) surface projected posteroventrally and facing dorsomedially (2), posteriorly elongated, triangular-shaped and facing dorsally (3), or posteroventrally projecting and paddleshaped (4). CHARACTER 72 (CLARK, 1994: character 72): Prearticular: present (0), or absent (1). CHARACTER 73 (modified from CLARK, 1994: character 73): + Articular without medial process (0), with short process not contacting braincase (1), or with process articulating with otoccipital and basisphenoid (2). CHARACTER 74 (CLARK, 1994: character 74): Dorsal edge of surangular: flat (0), or arched dorsally (1). CHARACTER 75 (CLARK, 1994: character 75): Mandibular fenestra: present (0), or absent (1). CHARACTER 76 (CLARK, 1994: character 76): Insertion area for M. pterygoideous posterior: does not extend onto lateral surface of angular (0), or extends onto lateral surface of angular (1). CHARACTER 77 (modified from CLARK, 1994: character 77): + Splenial involvement in symphysis in ventral view: not involved (0), involved slightly in symphysis (1), or extensively involved (2). CHARACTER 78 (CLARK, 1994: character 78): Posterior premaxillary teeth: similar in size to anterior teeth (0), or much longer (1). CHARACTER 79 (modified from CLARK, 1994: character 79): + Maxillary teeth waves: absent, no size variation (0), one wave of teeth enlarged (1), or enlarged maxillary teeth curved in two waves (“festooned”) (2). CHARACTER 80 (CLARK, 1994: character 80): Anterior dentary teeth opposite premaxilla-maxilla contact: no more than twice the length of other dentary teeth (0), or more than twice the length of other dentary teeth (1). CHARACTER 81 (modified from CLARK, 1994: character 81): Dentary teeth posterior to tooth opposite premaxilla- maxilla contact: equal in size (0), or enlarged dentary teeth opposite to smaller teeth in maxillary toothrow (1). CHARACTER 82 (modified from CLARK, 1994: character 82 by ORTEGA et al., 2000: character 120): Anterior and posterior scapular edges: symmetrical in lateral view (0), anterior edge more strongly concave than posterior edge (1), or dorsally narrow with straight edges (2).

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CHARACTER 83 (modified from CLARK, 1994: character 83 by ORTEGA et al., 2000: character 121): Coracoid length: up to two-thirds of the scapular length (0), or subequal in length to scapula (1). CHARACTER 84 (CLARK, 1994: character 84): Anterior process of ilium: similar in length to posterior process (0), or one-quarter or less of the length of the posterior process (1). CHARACTER 85 (CLARK, 1994: character 85): Pubis: rodlike without expanded distal end (0), or with expanded distal end (1). CHARACTER 86 (CLARK, 1994: character 86): Pubis: forms anterior half of ventral edge of acetabulum (0), or pubis at least partially excluded from the acetabulum by the anterior process of the ischium (1). CHARACTER 87 (CLARK, 1994: character 87): Distal end of femur: with large lateral facet for the fibula (0), or with very small facet (1). CHARACTER 88 (CLARK, 1994: character 88): Fifth pedal digit: with phalanges (0), or without phalanges (1). CHARACTER 89 (CLARK, 1994: character 89): intercentrum: broader than long (0), or as long as broad (1). CHARACTER 90 (modified from CLARK, 1994: character 90): + Cervical neural spines: all anteroposteriorly large (0), only posterior ones rodlike (1), or all spines rodlike (2). CHARACTER 91 (modified from CLARK, 1994: character 91 by BUSCALIONI & SANZ, 1988: character 37 and by BROCHU, 1997a: character 7): + Hypapophyses in cervicodorsal vertebrae: absent (0), present only in cervical vertebrae (1), present in cervical and the first two dorsal vertebrae (2), present up to the third dorsal vertebra (3), or present up to the fourth dorsal vertebrae (4). CHARACTER 92 (CLARK, 1994: character 92): Cervical vertebrae: amphicoelous or amphyplatian (0), or procoelous (1). CHARACTER 93 (CLARK, 1994: character 93): Trunk vertebrae: amphicoelous or amphyplatian (0), or procoelous (1). CHARACTER 94 (CLARK, 1994: character 94): All caudal vertebrae: amphicoelous or amphyplatian (0), first caudal biconvex with other procoelous (1), or procoelous (2). CHARACTER 95 (CLARK, 1994: character 95): Dorsal osteoderms: rounded or ovate (0), or rectangular, broader than long (1), or square (2). CHARACTER 96 (modified from CLARK, 1994: character 96, and BROCHU, 1997a: character 40): + Dorsal osteoderms: without articular anterior process (0), with a discrete convexity on anterior margin (1), or with a well-developed process located anterolaterally in dorsal parasagittal osteoderms (2). CHARACTER 97 (modified from CLARK, 1994: character 97 by ORTEGA et al., 2000: characters. 107 and 108): + Rows of dorsal osteoderms: two parallel rows (0), more than two rows (1), or more than four rows with “accessory ranges of osteoderms” (sensu Frey, 1988) (2). CHARACTER 98 (CLARK, 1994: character 98): Osteoderms: some or all imbricated (0), or sutured to one another (1). CHARACTER 99 (CLARK, 1994: character 99): Tail osteoderms: dorsal only (0), or completely surrounded by osteoderms (1). CHARACTER 100 (CLARK, 1994: character 100): Trunk osteoderms: absent from ventral part of the trunk (0), or present (1). CHARACTER 101 (CLARK, 1994: character 101): Osteoderms: with longitudinal keels on dorsal surfaces (0), or without longitudinal keels (1). CHARACTER 102 (WU & SUES, 1996: character 14): Jugal: participating in margin of antorbital fossa (0), or separated from it (1). CHARACTER 103 (modified from WU & SUES, 1996: character 23): + Articular facet for quadrate condyle: equal in length to the quadrate condyles (0), slightly longer (1), or close to three times the length of the quadrate condyles (2). CHARACTER 104 (modified from WU & SUES, 1996: character 24 and WU et al., 1997: character 124): + Jaw joint: placed at level with basioccipital condyle (0), below basioccipital condyle about above level of lower toothrow (1), or below level of toothrow (2). CHARACTER 105 (modified from WU & SUES, 1996: character 27 and ORTEGA et al., 2000: character 133): + Premaxillary teeth: five (0), four (1), three (2), or two (3). CHARACTER 106 (modified from WU & SUES, 1996: character 29): Unsculptured region along alveolar margin on lateral surface of maxilla: absent (0), or present (1). CHARACTER 107 (WU & SUES, 1996: character 30): + Maxilla: with eight or more teeth (0), seven teeth (1), six teeth (2), five teeth (3), or four teeth (4). CHARACTER 108 (WU & SUES, 1996: character 33): Coracoid: without posteromedial or ventromedial process (0), with elongate posteromedial process (1), or distally expanded ventromedial process (2). CHARACTER 109 (WU & SUES, 1996: character 40): Radiale and ulnare: short and massive (0), or elongate (1). CHARACTER 110 (WU & SUES, 1996: character 41): Postacetabular process: directed posteroventrally or posteriorly (0), or directed posterodorsally and much higher in position than preacetabular process (1). CHARACTER 111 (modified from GOMANI, 1997: character 4): Prefrontals anterior to orbits: elongated, oriented parallel to anteroposterior axis of the skull (0), or short and broad, oriented posteromedially-anterolaterally (1). CHARACTER 112 (modified from GOMANI, 1997: character 32): Basioccipital and ventral part of otoccipital: facing posteriorly (0), or facing posteroventrally (1). CHARACTER 113 (BUSCALIONI & SANZ, 1988: character 35): Vertebral centra: cylindrical (0), or spool shaped (1). CHARACTER 114 (modified from BUSCALIONI & SANZ, 1988: character 39): Transverse process of posterior dorsal vertebrae

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dorsoventrally low and laminar (0), or dorsoventrally high (1). CHARACTER 115 (BUSCALIONI & SANZ, 1988: character 44): Number of sacral vertebrae: two (0), or more than two (1). CHARACTER 116 (BUSCALIONI & SANZ, 1988: character 49): Supra-acetabular crest: present (0), or absent (1). CHARACTER 117 (BUSCALIONI & SANZ, 1988: character 54): Proximal end of radiale expanded symmetrically, similarly to the distal end (0), or more expanded proximomedially than proximolaterally (1). CHARACTER 118 (ORTEGA et al., 1996: character 5): Lateral surface of the dentary: without a longitudinal depression (0), or with a longitudinal depression (1). CHARACTER 119 (ORTEGA et al., 1996: character 9): Ventral exposure of splenials: absent (0), or present (1). CHARACTER 120 (ORTEGA et al., 1996: character 11, ORTEGA et al., 2000: character 100): Tooth margins: with denticulate carinae (0), or without carinae or with smooth or crenulated carinae (1). CHARACTER 121 (modified from POL, 1999a: character 133 and ORTEGA et al., 2000: character 145): Lateral surface of anterior process of jugal: flat or convex (0), or with broad shelf below the orbit with triangular depression underneath it (1). CHARACTER 122 (POL, 1999a: character 134): Jugal: does not exceed the anterior margin of orbit (0), or exceeds the anterior margin of orbit (1). CHARACTER 123 (POL, 1999a: character 135): Notch in premaxilla on lateral edge of external nares: absent (0), or present on the dorsal half of the external nares lateral margin (1). CHARACTER 124 (POL, 1999a: character 136): Dorsal border of external nares: formed mostly by the nasals (0), or by both the nasals and premaxilla (1). CHARACTER 125 (POL, 1999a: character 138): Posterodorsal process of premaxilla: absent (0), or present extending posteriorly wedging between maxilla and nasals (1). CHARACTER 126 (POL, 1999a: character 139 and ORTEGA et al., 2000: character 9): + Premaxilla-maxilla suture in palatal view, medial to alveolar region: anteromedially directed (0), sinusoidal, posteromedially directed on its lateral half and anteromedially directed along its medial region (1), or posteromedially directed (2). CHARACTER 127 (POL, 1999a: character 140): Nasal lateral border posterior to external nares: laterally concave (0), or straight (1). CHARACTER 128 (POL, 1999a: character 141): Nasal lateral edges: nearly parallel (0), oblique to each other converging anteriorly (1), or oblique to each other diverging anteriorly (2). CHARACTER 129 (POL, 1999a: character 143): Palatine anteromedial margin: exceeding the anterior margin of the palatal fenestrae wedging between the maxillae (0), or not exceeding the anterior margin of palatal fenestrae (1). CHARACTER 130 (POL, 1999a: character 144): Dorsoventral height of jugal antorbital region respect to infraorbital region: equal or lower (0), or antorbital region more expanded than infraorbital region of jugal (1). CHARACTER 131 (POL, 1999a: character 145): Maxilla-lacrimal contact: partially included in antorbital fossa (0), or completely included in antorbital fossa (1). CHARACTER 132 (POL, 1999a: character 146): Lateral eustachian tube openings: located posteriorly to the medial opening (0), or aligned anteroposteriorly and dorsoventrally (1). CHARACTER 133 (POL, 1999a: character 147): Anterior process of ectopterygoid: developed (0), or reduced–absent (1). CHARACTER 134 (POL, 1999a: character 148): Posterior process of ectopterygoid: developed (0), or reduced-absent (1). CHARACTER 135 (POL, 1999a: character 149 and ORTEGA et al., 2000: character 13): Small foramen located in the premaxillo-maxillary suture in lateral surface (not for big mandibular teeth): absent (0), or present (1). CHARACTER 136 (POL, 1999a: character 150): Jugal posterior process: exceeding posteriorly the infratemporal fenestrae (0), or not (1). CHARACTER 137 (POL, 1999a: character 151): Compressed crown of maxillary teeth: oriented parallel to the longitudinal axis of skull (0), or obliquely disposed (1). CHARACTER 138 (POL, 1999a: character 152): Large and aligned neurovascular foramina on lateral maxilary surface: absent (0), or present (1). CHARACTER 139 (modified from POL, 1999a: character 153): External surface of maxilla and premaxilla: with a single plane facing laterally (0), or with ventral region facing laterally and dorsal region facing dorsolaterally (1). CHARACTER 140 (POL, 1999a: character 154 and ORTEGA et al., 2000: character 104): Maxillary teeth: not compressed laterally (0), or compressed laterally (1). CHARACTER 141 (POL, 1999a: character 155): Posteroventral corner of quadratojugal: reaching the quadrate condyles (0), or not reaching the quadrate condyles (1). CHARACTER 142 (POL, 1999a: character 156): Base of postorbital process of jugal: directed posterodorsally (0), or directed dorsally (1). CHARACTER 143 (POL, 1999a: character 157): + Postorbital process of jugal: anteriorly placed (0), in the middle (1), or posteriorly positioned (2). CHARACTER 144 (POL, 1999a: character 158 and ORTEGA et al., 2000: character 36): Postorbital-ectopterygoid contact: present (0), or absent (1). CHARACTER 145 (POL, 1999a: character 161): Quadratojugal: not ornamented (0), or ornamented in the base (1).

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CHARACTER 146 (POL, 1999a: character 162): Prefrontal-maxillary contact in the inner anteromedial region of orbit: absent (0), or present (1). CHARACTER 147 (POL, 1999a: character 163): Basisphenoid: without lateral exposure (0), or with lateral exposure on the braincase (1). CHARACTER 148 (POL, 1999a: character 165): Quadrate process of pterygoids: well developed (0), or poorly developed (1). CHARACTER 149 (modified from POL, 1999a: character 166 and ORTEGA et al., 2000: character 44): + Quadrate major axis directed: posteroventrally (0), ventrally (1), or anteroventrally (2). CHARACTER 150 (POL, 1999a: character 167): Quadrate distal end: with only one plane facing posteriorly (0), or with two distinct faces in posterior view, a posterior one and a medial one bearing the foramen aereum (1). CHARACTER 151 (POL, 1999a: character 168): Anteroposterior development of neural spine in axis: well developed covering all the neural arch length (0), or poorly developed, located over the posterior half of the neural arch (1). CHARACTER 152 (POL, 1999a: character 169): Prezygapophyses of axis: not exceeding anterior edge of neural arch (0), or exceeding the anterior margin of neural arch (1). CHARACTER 153 (POL, 1999a: character 170): Postzygapophyses of axis: well developed, curved laterally (0), or poorly developed (1). CHARACTER 154 (modified from POL, 1999b: character 212): Shape of dentary symphysis in ventral view: tapering anteriorly forming an angle (0), Ushaped, smoothly curving anteriorly (1), or lateral edges longitudinally oriented, convex anterolateral corner, and extensive transversally oriented anterior edge (2). CHARACTER 155 (POL, 1999b: character 213): Unsculpted region in the dentary below the tooth row: absent (0), or present (1). CHARACTER 156 (ORTEGA et al., 1996: character 13 and BUCKLEY et al., 2000: character 117): Cheek teeth: not constricted at base of crown (0), or constricted at base of crown (1). CHARACTER 157 (ORTEGA et al., 2000: character 42): Outer surface of squamosal laterodorsally oriented: extensive (0), or reduced and sculpted (1), or reduced and unsculpted (2). CHARACTER 158 (ORTEGA et al., 2000: character 74): Length/height proportion of infratemporal fenestra: higher than long or subequal (0), or very anteroposteriorly elongated (1). CHARACTER 159 (ORTEGA et al., 2000: character 90): Foramen intramandibularis oralis: small or absent (0), or big and slotlike (1). CHARACTER 160 (ORTEGA et al., 2000: character 146): Ectopterygoid medial process: single (0), or forked (1). CHARACTER 161 (modified from GOMANI, 1997: character 46 and BUCKLEY et al., 2000: character 113): Cusps of teeth: unique cusp (0), one main cusp with smaller cusps arranged in one row (1), one main cusp with smaller cusps arranged in more than one row (2), several cusps of equal size arranged in more than one row (3), or multiple small cusps along edges of occlusal surface (4). CHARACTER 162 (POL & NORELL, 2004a: character 164): Cross section of distal end of quadrate: mediolaterally wide and anteroposteriorly thin (0), or subquadrangular (1). CHARACTER 163 (POL & NORELL, 2004a: character 165): Palatine-pterygoid contact on palate: palatines overlie pterygoids (0), or palatines firmly sutured to pterygoids (1). CHARACTER 164 (WU et al., 1997: character 103): Squamosal descending process: absent (0), or present (1). CHARACTER 165 (modified from WU et al., 1997: character 105): + Development of distal quadrate body ventral to otoccipital-quadrate contact: distinct (0), incipiently distinct (1), or indistinct (2). CHARACTER 166 (WU et al., 1997: character 106): Pterygoid flanges: thin and laminar (0), or dorsoventrally thick, with pneumatic spaces (1). CHARACTER 167 (WU et al., 1997: character 108): Postorbital participation in infratemporal fenestra: almost or entirely excluded (0), or bordering infratemporal fenestra (1). CHARACTER 168 (WU et al., 1997: character 109): Palatines: form margin of suborbital fenestra (0), or excluded from margin of suborbital fenestra (1). CHARACTER 169 (WU et al., 1997: character 110): Angular posterior to mandibular fenestra: widely exposed on lateral surface of (0), or shifted to the ventral surface of mandible (1). CHARACTER 170 (WU et al., 1997: character 112): Posteroventral edge of mandibular ramus: straight or convex (0), or markedly deflected (1). CHARACTER 171 (modified from WU et al., 1997: character 119): Quadrate ramus of pterygoid in ventral view: narrow (0), or broad (1). CHARACTER 172 (WU et al., 1997: character 121): Pterygoids: not in contact anterior to basisphenoid on palate (0), or pterygoids in contact (1). CHARACTER 173 (WU et al., 1997: character 122): Olecranon: well developed (0), or absent (1). CHARACTER 174 (WU et al., 1997: character 123): Cranial table width respect to ventral portion of skull: as wide as ventral portion of skull (0), or narrower than ventral portion of skull (1). CHARACTER 175 (WU et al., 1997: character 127): Depression on posterolateral surface of maxilla: absent (0), or present (1).

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CHARACTER 176 (WU et al., 1997: character 128): Anterior palatal fenestra: absent (0), or present (1). CHARACTER 177 (POL & NORELL, 2004a: character 179): Paired ridges located medially on ventral surface of basisphenoid: absent (0), or present (1). CHARACTER 178 (POL & NORELL, 2004a: character 180): Posterolateral end of quadratojugal: acute or rounded, tightly overlapping the quadrate (0), or with sinusoidal ventral edge and wide and rounded posterior edge slightly overhanging the lateral surface of the quadrate (1). CHARACTER 179 (POL & NORELL, 2004a: character 181): Orientation of quadrate body distal to otoccipital-quadrate contact in posterior view: ventrally (0), or ventrolaterally (1). CHARACTER 180 (GASPARINI et al., 1993: character 3): Wedgelike process of the maxilla in lateral surface of premaxilla- maxilla suture: absent (0), or present (1). CHARACTER 181 (POL & NORELL, 2004b: character 181): Palpebrals: separated from the lateral edge of the frontals (0), or extensively sutured to each other and to the lateral margin of the frontals (1). CHARACTER 182 (POL & NORELL, 2004b: character 182): External surface of ascending process of jugal: exposed laterally (0), or exposed posterolaterally (1). CHARACTER 183 (POL & NORELL, 2004b: character183): Longitudinal ridge on lateral surface of jugal below infratemporal fenestra: absent (0), or present (1). CHARACTER 184 (POL & NORELL, 2004b: character 184): Dorsal surface of posterolateral region of squamosal: without ridges (0), or with three curved ridges oriented longitudinally (1). CHARACTER 185 (POL & NORELL, 2004b: character 185): Ridge along dorsal section of quadrate-quadratojugal contact: absent (0), or present (1). CHARACTER 186 (POL & NORELL, 2004b: character 186): Sharp ridge along the ventral surface of angular: absent (0), or present (1). CHARACTER 187 (POL & NORELL, 2004b: character 187): Longitudinal ridge along the dorsolateral surface of surangular: absent (0), or present (1). CHARACTER 188 (POL & NORELL, 2004b: character 188): Dorsal surface of osteoderms ornamented with anterolaterally and anteromedially directed ridges (fleur de lys pattern of OSMÓLSKA et al., 1997): absent (0), or present (1). CHARACTER 189 (POL & NORELL, 2004b: character 189): Cervical region surrounded by lateral and ventral osteoderms sutured to the dorsal elements: absent (0), or present (1). CHARACTER 190 (POL & NORELL, 2004b: character 190): Appendicular osteoderms: absent (0), or present (1). CHARACTER 191 (ORTEGA et al., 2000: character 72): Supratemporal fenestra: present (0), or absent (1). CHARACTER 192 (POL & NORELL, 2004a: character 183): Choanal opening: opened posteriorly and continuous with pterygoid surface (0), or closed posteriorly by an elevated wall formed by the pterygoids (1). CHARACTER 193 (POL et al., 2004: caract. 164) Major axis of ectopterygoid body oriented: anterolaterally (0), or anteriorly (1). CHARACTER 194 (POL et al., 2004: character 179): Ventral margin of infratemporal bar of jugal: straight (0), or dorsally arched (1). CHARACTER 195 (WU & SUES, 1996: character 6): Premaxilla-maxilla segment longer than (0) or shorter than (1) remainder of skull in lateral view. CHARACTER 196 (WU & SUES, 1996: character 17): deep (0) or shallow and spatulate anteriorly (1). CHARACTER 197 (WU & SUES, 1996: character 31): Maxillary tooth row extending posterior to anterior border of orbit (0) or terminating in front of orbit (1) in lateral view. CHARACTER 198 (MARTINELLI, 2003: character 35): Ectopterygoid does not contact posterior part of palatine (0), or contact palatine, excluding the pterygoid of the posterior edge of the fenestra palatina (1). CHARACTER 199 (MARTINELLI, 2003: character 36): Nasal-frontal suture transversely oriented (0) or obliquely oriented (1). CHARACTER 200 (FIORELLI, 2005): Hipapophysis in cervical vertebrae: absent (0), like a vertical thorn slightly or well marked (1) or like keel-shaped running anteroposteriorly in ventral surface of centrum (2). CHARACTER 201 (FIORELLI, 2005): First and second pair of mandibular teeth directed, in relation to the vertical one, toward up practically vertical (0) or directed anterodorsally in an angle approximate of 45º-50º (1). CHARACTER 202 (FIORELLI, 2005): Postcanines teeth (molariforms) triangular in transverse section (0), rounded, cuspidate or tablets laterally (Ziphodont or basal type) (1). CHARACTER 203 (FIORELLI, 2005): Small and big neurovascular foramina aligned on lateral surface of dentary: absent (0) or present (1). CHARACTER 204 (FIORELLI, 2005): Anteroposterior crest directed in the glenoid fossa on articular shelf separating the articulation cavities for the respective condyles of quadrate: absent (0) or present (1). CHARACTER 205 (FIORELLI, 2005): Posterior Buttress on shelf of articular like top for the quadrate: absent (0) or present (1). CHARACTER 206 (FIORELLI, 2005): Rounded cervical centra (0) in transverse section or irregulary polygonal (heptagonal) formed one of their vertexes the ventral keel (hipapophysis) (1). CHARACTER 207 (FIORELLI, 2005): Development of thin pre and postspinals sheets in anterior dorsal vertebrae: absent or little developed (0) or developed (1).

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CHARACTER 208 (FIORELLI, 2005): Suprapostzygapophyseal laminae in cervical and cervicodorsal vertebrae: absent (0) or present (1). CHARACTER 209 (FIORELLI, 2005): Development of the acetabular roof of ilium with a deep acetabular cavity: not developed (0) or well developed (1). CHARACTER 210 (FIORELLI, 2005): Prominent process on femur (for m. coccygeofemoralis) located medially in the proximal end of shaft: absent or slightly developed (0) or very developed (1). CHARACTER 211 Cervical vertebrae centra very anteroposteriorly lengthened (0), or shorter and tablets in anteroposterior sense (1). CHARACTER 212 Articulation surface of the parapophysis for the chapter of the ribs in cervical vertebrae: anteroposteriorly lengthened –double long than wide or more– (0) or practically square or rounded – as long as wide – (1). CHARACTER 213 Long postparapophyseal border in cervical vertebrae, anteroposteriorly directed until the posterior border of the centrum, forming deep furrows toward both sides (up and below) of the parapophyseal border: absent (0), present (1). CHARACTER 214 Hook or expansion in the posterior vertex of the scapula formed by the posterior and dorsal border: absent (0), present (1). CHARACTER 215 (POL, 1999a: character192) Lateral expansion in proximal extreme of the humerus: absent (0), present (1). CHARACTER 216 Proportion among the long of the deltopectoral crest (Dc) in relation to the total length (TL) of the humerus (= Dc hu / TL hu): smaller than 25 % (0) or bigger than 25 % (1). CHARACTER 217 Proportion among the diameter of the shaft (Dsh) of the humerus measured in half of their longitude in relation to the total length (TL) of the humerus (= Dsh hu/ TL hu): smaller or similar to 7 % (0) or bigger than 7 % (1). CHARACTER 218 (modified from POL, 1999a: character 191) + Proportion among the total length of humerus and wide of proximal expansion: in the range between 2.15 and 2.3 (0), between 2.8 and 3.2 (1), bigger at 3.7 and 4.74 (2), same or bigger at 5.0 (3). CHARACTER 219 Proportion among the diameter of the shaft (Dsh ra) of the radius measured in half of their longitude in relation to the total length (TL ra) of the radius (= Dsh ra / TL ra): smaller or similar to 4 % (0), between 4 % and 6 % (1) or bigger than 6 % (2). CHARACTER 220 Relationship between the total length of the ulna and the total length of the humerus (= TL ul / TL hu): ulna < humerus (0) or ulna > humerus (1). CHARACTER 221 Relationship between the broad of the shaft of ulna and their total length (= BS ul / TL ul): smaller than 5 % (0), between 5 % and 7 % (1) or bigger than 7 % (2). CHARACTER 222 Broad of the femoral shaft in relation to their total length (= BS fe / TL fe): smaller than 9 % (0) or bigger than 9 % (1). CHARACTER 223 Broad of the tibial shaft in relation to their total length (= BS ti / TL ti): smaller or similar to 7 % (0) or bigger than 7 % (1). CHARACTER 224 Relationship among the broad of the distal expansion of pubis (B.d.e pu) and the total length (TL pu) of the same one (= B.d.e pu / TL pu): smaller or similar to 30 % (0) or bigger than 30 % (1). CHARACTER 225 Relationship among the diameter of the pubic shaft (D.sh.pu) and the total length (TL pu) of the same one (= D.sh.pu / TL pu): smaller than 8 % (0) or bigger than 8 % (1). CHARACTER 226 (modified from SERENO, 1991: character 27): Relationship between the total length of the femur and the total length of the tibia (= TL fe / TL ti): femur > tibia (0) or femur < tibia (1). CHARACTER 227 Anteroposterior longitudinal relationship between the ventral scapular section (v.S) and dorsal scapular blade (d.S) [= v.S/d.S]: smaller than 55 % (0); between 55 % and 70 % (1); between 70 % and 100 % (2) or bigger than 100 % (3). CHARACTER 228 Relationship between the anteroposterior length of dorsal scapular blade (d.S) and the major dorsoventral longitudinal axis (m.l.a.S) of the same one [= d.S/m.l.a.S]: smaller than 40 % (0); between 40 % and 55 % (1) or bigger than 55 % (2). CHARACTER 229 Relationship between the diameter of the scapular half constriction (S.h.c) and the major dorsoventral longitudinal axis (m.l.a.S) of the same one [= S.h.c/ m.l.a.S]: less than 15 % (0); between 15 % and 20 % (1) or more than 20 % (2). CHARACTER 230 Relationship between the major dorsoventral longitudinal axis of scapula (m.l.a.S) and the total length of the humerus (t.l.hu) [= m.l.a.S/t.l.hu]: less than 70 % (0) or more than 70 % (1). CHARACTER 231 Relationship between the total length of the pubis (t.l.pu) and the total length of femur (t.l.fe) [= t.l.pu/t.l.fe]: less than 45 % (0) or more than 45 % (1).

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APPENDIX II

List of the 51 taxa used in the phylogenetic analysis (taken from POL & NORELL, 2004b; POL et al., 2004). , Mariliasuchus, Candidodon, , and Uberabasuchus are new taxa included in this paper.

Gracilisuchus stipanicicorum (ROMER, 1972) Terrestrisuchus gracilis (CRUSH, 1984) Dibothrosuchus elaphros (WU & CHATTERJEE, 1993) Protosuchus richardsoni (COLBERT & MOOK, 1951) Hemiprotosuchus leali (BONAPARTE, 1971) Kayenta Form (CLARK, 1986) Edentosuchus tienshanensis (YOUNG, 1973; POL et al., 2004) Orthosuchus stormbergi (NASH, 1975) Gobiosuchus kielanae (OSMÓLSKA, 1972) Zaraasuchus shepardi (POL & NORELL, 2004b) Shantungosuchus hangjinensis (WU et al., 1994) Neuquensuchus universitas (MUCPv-47, MUCPv-161) Sichuanosuchus shuhanensis (WU et al., 1997) Zosuchus davidsoni (POL & NORELL, 2004a) Fruita Form (CLARK, 1985, 1994) Hsisosuchus chungkingensis (YOUNG & CHOW, 1953; LI et al., 1994; WU et al., 1994) Notosuchus terrestris (WOODWARD, 1896; GASPARINI, 1971) Anatosuchus minor (SERENO et al., 2003) Comahuesuchus brachybuccalis (BONAPARTE, 1991) Mariliasuchus amarali (CARVALHO & BERTINI, 1999) Uruguaysuchus aznarezi (RUSCONI, 1933) Chimaeresuchus paradoxus (WU & SUES, 1996) mwakasyungutiensis (CLARK et al., 1989; GOMANI, 1997) Candidodon itapecuruense (CARVALHO, 1994; NOBRE & CARVALHO, 2002) Simosuchus clarki (BUCKLEY et al., 2000) Sphagesaurus huenei (PRICE, 1950; POL, 2003) Araripesuchus gomesii (PRICE, 1959) Araripesuchus patagonicus (ORTEGA et al., 2000) pachecoi (PRICE, 1945) Stratiotosuchus maxhechti (CAMPOS et al., 2001) Bretesuchus bonapartei (GASPARINI et al., 1993) Iberosuchus macrodon (ANTUNES, 1975; ORTEGA et al., 2000) Lomasuchus palpebrosus (GASPARINI et al., 1991) torminni (PRICE, 1955; GASPARINI et al., 1991) Uberabasuchus terrificus (CARVALHO et al., 2004) Theriosuchus pusillus (OWEN, 1879; CLARK, 1986, 1994; ORTEGA et al., 2000) Alligatorium (WELLNHOFER, 1971; CLARK, 1986, 1994) Eutretauranosuchus delfsi (MOOK, 1967; CLARK, 1986, 1994) Goniopholis (MOOK, 1942; CLARK, 1986, 1994; SALISBURY et al., 1999) decipiens (OWEN, 1878; CLARK, 1986, 1994) Dyrosaurus phosphaticus (BUFFETAUT, 1978; CLARK, 1986, 1994) Sokotosuchus ianwilsoni (HALSTEAD, 1975; BUFFETAUT, 1979; CLARK, 1986, 1994) Pelagosaurus typus (EUDES-DESLONGCHAMPS, 1863) (BUFFETAUT, 1982; CLARK, 1986, 1994) Metriorhynchidae (KÄLIN, 1955; GASPARINI & DIAZ, 1977) Hylaeochampsa vectiana (CLARK & NORELL, 1992; ORTEGA et al., 2000) Bernissartia fagessi (BUSCALIONI & SANZ, 1990; NORELL & CLARK, 1990) formidabilis (ERICKSON, 1976; BROCHU, 1997b) Gavialis gangeticus (CLARK, 1994; BROCHU, 1997a) Crocodylus niloticus (CLARK, 1994; BROCHU, 1997a) Alligator mississippiensis (CLARK, 1994; BROCHU, 1997a)

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APPENDIX III

DATA MATRIX USED IN PHYLOGENETIC ANALYSIS

Gracilisuchus stipanicicorum 000000??0?000000000000?0?000000000?0??0?0?00000?000???0000?0???00000?100000?00000000?011?0000? 0?000001012?00?00???00?01?01000??1?01???000001002?0???0000???0?0??0??0000?0000000?00000000?0?00 000?00000000100?000?00100111320201??03011?

Terrestrisuchus gracilis 000??00??01?000000?000001000000?110?00000?00000?000??0?000?000????002??0100?000000000010?0000? 02000001010?01100?1000000001001?10?00?11010001?[01]110???00000??0000000?0?0??00??0100??00??000? ????00000010100100?0001001100003010000?121001

Dibothrosuchus elaphros 000?00?020??001???000000??????00110000000?00000?0000?00000?0?0101000?010100?0010?000?????2000? 0?????01010?01100?0?00000001001?10?00?1?000101011100??000001??00000010001000100?0?00000000?00 0000000001?100100??00100???1??2?110?????????

Protosuchus richardsoni 2100000120?000011010002100000100010001010?00201001111110010101102011?110210001010100011100[1 234]00?120011010111021001010000[01]000000?01??01??10010[01]0101000000???0100000000120000011110? ?01000?010?000000000001201100???10111011122011100030011

Hemiprotosuchus leali ?00?00?10??????10010?0??00?0010?11?0??01??0020?00?11?1100101??1?2?11??1?21????01?????????0????120 0?1?101??0??????????000?000??10?00???00000??10?????00???????0?000?12???001??10?0?00?01??00???0000 00000?01?00?????????0????????????????

Kayenta Form [12]01110?1200000?10010?0??00????0?0???11110?002010011111100001011?2011?0102100?1010??0????00?? 0?1200101101112?????0????01100?00?01000111?101001?01?10000000011?0??40012??00011??0?00???00???? ????011100???01?????????????????????????????

Edentosuchus tienshanensis 201?????[12]????0??[01]0??1?0100??0?????02?110?00?????????????????????[12]?311????10?01010?????????[23 4]??????????????1[23]??????????00110??1?01?1???1000110?11?1????????011????4?0??1????11??0????0100??? ????00111000??01100???10110??????????????????

Orthosuchus stormbergi 21100001201?0001001000[01]10000010001000?000?002011001111100??1?1?02011?0?0?0?001000100011100 000?120010010211421001?10010?100000001?01010000000000?0???00001???000000?12?00001111000010000 0?000?0000010001?01?00???10?1?00??3?0?1???0?????

Gobiosuchus kielanae 101000?110000011001?[01][01]?1?00001?10?0201000?0020112011111000?1????201???1?20100[01]010?0?0??? ????0?1010110[01]012002??0000???0010[01]00001000000?00001001211?0000???110000000?121000011?00?0 ?0011111111111000100?1?01?0????100??0000??0100000???00

Zaraasuchus shepardi 10?????????????1?01?01?1000001?10?02????????????????????????????2?????1??010??????????????[1234]0??10 10??0???????????0??????????????????????0?????1??1?????????1?00????????1?00???0??????11111111111?00?? ??00???000????001???0???1??????????

Shantungosuchus hangjinensis 2?1????1?0???0?1??1????11??????????21?1[01]100020?1?011?1100?10????????101?1?000??10???????0?1????? ???????1???????1?????0010???????00??10?00??111211??001?????0?0?00????10111111?0?110??0???1?????00[0 1]1000?201??0???10001??00??0100001????0

Neuquensuchus universitas ?????????????????????????????????????????????????????????????????????????????????1??0111?11000????????? ??????0??1?0?????????????????????????????????????????????????????????0???????????????00?????????2?????0 ???0001110030110000102100

Sichuanosuchus shuhanensis [12]01??0?1200[01]00?10010[01]1?110???1?00?021?10?00020?1?011?1100???????2?11????1?000011?1??01???? 000????????1?11?0?1????0??100100??1??10?0????00111[01]1210??00?????1?????010111011111100?11000010 0?1???00?010001201000?00?0?0111002001??00?0210?

Arq. Mus. Nac., Rio de Janeiro, v.65, n.4, p.417-459, out./dez.2007 FIRST CRETACEOUS “PROTOSUCHIAN” FROM GONDWANA 455

Zosuchus davidsoni 201??0?1200000??001010[01]110?001110?02211010012?1??011?11000?0?1?0211110????0?01111???????????? ????????1?12?3????1?????00100011011?0001?0?0010112?[01]?0001???0?00???010111??1011?1011100000010 0???000010001?011?0??????????????????????????

Fruita Form 201??001200100010000100100000110010221?11?0020112?1???0?0??0??1?2?31?????1?0111101011?1?000111 12?0??1???[01]00???1?1001?001?0?0100100??101?0011?01110??0??00?10?0000?1???000????101?0?00000?00 0??0?0??0?????1?????????????????????????????

Hsisosuchus chungkingensis 211??????1??000000100001100011000?0221101000[12]??12?11?10000?0?1?0??111?4?00[01]02?1??10???????0 00?1000???101?0021??1?????01001???????0000??00??1?11?1??00????????0??0????10?0?0111[01]?00??00?0?1 000?01?0001012?1?0???????????????02?????2121?

Notosuchus terrestris 101?001101010011100011111100110011022110110021112011?1000010?110211111210101110001[01]11101? 210001000??0122011??01100101[01]1101[01]010010000001111111011000001110010000101110110000111011 0000000000000010011111210110111111100111120211??01221?

Mariliasuchus amarali 101?00?1?10?00?11000101111?0111?01022?10???021?120?1?100??1??1??2130?121010111000??1??0??2?000? ???????2201???011?010?01101101001?0?000111111101?0??001???010000?0111?11?0001?101?0?00000000?0 00100111?1?10110?11?1?????????0211??01221?

Anatosuchus minor 203?00?1?11?001?1?00101111?0010101022?1010?01??????????0?????????1212131?10?10101???????????????? ????1?1000???0??????01101?01?10?00????00?00111?00??0????10020??0111?1?000???100???0?000000???010 001000??1?00??????????????????????????

Comahuesuchus brachybuccalis 103??0?1011?00?????0112????????0010?2????1??11?1?????????????????131??????0?10101??????????????????? ????[01]13???1??????0?10?101201?01?????011??0?1????11???11?00100?1?0??000???100??0?000??0????0?0?1 1?11?111????????????????????????????

Sphagesaurus huenei 101?000101??00??100?????110?????????21101?00?????011?1000????????13?2????????100????????1?????????? ?????312????0???????11111101111111111111110011101011?0?11?0??011?0?10??01??000000?00???????0100 1?1????0?????????????????????????????

Chimaerasuchus paradoxus 101?0001111?00???????????????????????????????????????????????????12??0110?01010??1?1?????2100?00???? 11[12]?314210??00?0100111111011??????0?0110??????????10?11?????3???????????1?00???0??????????0???11 1?1211?101111?110011?1202????01221?

Malawisuchus mwakasyungutiensis 101?00?1110000?[01]10001[01][01]1100?110001?22110100011??20???1000?10?1?02?111[01]2?0101110001???? 1??210000010??01[12]2111???01?0???01100101?11000???110110101?0?0001???0?100??21110?100001110000 000000000???0100110002?1110101??11001112?0211??01221?

Uruguaysuchus aznarezi 201?001101??00??10??1??1????1???01022?101?0011????1?????0??0???01111[12]???000110100??1?1??????000 0?0??01?21002100?00?000?[01]???01?1?00????1?0111?11?????11?????1?0001????????0???10?????00????????0 1?001101?01100?11?11?001111?????????????

Candidodon itapecuruense 201?00???1??00??110010?????0010101022?111?00????????????????????11212????????11???????????????????? ????1?01????????????1011010??00????100111111????11??????1?0??21?1?1???????1?0???00?000?????01101?0 00??1?????????????????????????????

Simosuchus clarki 10301011000000100010111110?0110001021?10100011?11011?1000010?1?020112121010110000???????02100 ?2010?10002010???01??????11011012120000101001110021100120???211[12]0001111011001[01]1?100000000 00010000010110100001100?????110?????202??????????

Araripesuchus gomesii 201000110100001110001011111011[01]001022110100011112011?10000?0?110201121210001101[01][01]1[01] 11111?1[234]000100010011110021001001010100100100100000010011000210000110?0011[01]000011110100 0011100?0000000000000010010000?01?0??????110011121021111012211

Arq. Mus. Nac., Rio de Janeiro, v.65, n.4, p.417-459, out./dez.2007 456 L.E.FIORELLI & J.O.CALVO

Araripesuchus patagonicus 201000?1010000?1[01]000101111?0111001022?10100011?12?11?1000??0?1?02?11212?0?011[01]1??1?1?????? ????1000??0111100???01???0?01?01101?010000??100110102?0??01????0??[01]1000111?0100001110?0000000 0000000010010000?01100?1?111100111210211??01221?

Baurusuchus pachecoi 100??0?121??00?1101????111?0110?????2?10110011112011?1000?10??10??311121010111111??????????????? ??????12103????1?????1101110101011100110011110110?0111???[01]0[01]1111011101?00001?0001000000000 0???010000101?01000??????????????????????????

Stratiotosuchus maxhechti 100?0??1??0000?11?111101?0???10?11??????????1?1??????????????????130?????????11????????????????????? ?0?2103???1??????????1111?10?1????010001101?10??1??????010??01????1??????10??010?0000????00??0??1? 1??1?????????????????????????????

Bretesuchus bonapartei 1[01]0??01121??00???????????0??????????2???10011????????1011?1??????13?1??1?00?10110???????????????? ???????100????1???????01??0????01??0???0??1?0???????????[01]0[01]?1?10?1???1??001??00??????????0????1? ?00101?0100??????????????????????

Iberosuchus macrodon 1?0?00012?0?00111000111111?01?000?02??10100111?12??1?101??10?1????111??10?0?1011011??????[12][12 34]00??00???00?[12][01]0?2??0000???11001101010?1?0??100?11001?0??101???[01]?0111?001101?0??01?1000 01000000??0??010?00????0100???????????????????????????

Lomasuchus palpebrosus 201????1211?00?11000101111??110001022?1010001??12??1?100??1??1??2?21?????00??0[12]11????????????? ???????1???00???00?????0?00???1?110?00???00011?0??1??0??????010???0?11??10??01?1000??11000??0???01 ?000100??1?????????????????????????????

Peirosaurus torminni 201?011??1??00??????10?1????????0???2?10???????????????????????????1??????????[12]1??????????????????? ?????000????????????0????1???0???????0?1??????????????[01]??????0??????????????00??1??????????0???00?00 ??1?????????????????????????????

Uberabasuchus terrificus 201100?1211?00?1100010?1?1?0110101122??0?0001????????????????????111?131110?11[12]11?????????????? ??????011000???0??????01101?01?10?00?1?010000101?11??0????10010??00?1??1000???100?0?1?000000???0 ?1000100?01001??????????????????????????

Theriosuchus pusillus 20110111110100110000110111100110011?211010001?11?01111000?????1?20211?4100101010110111110001 1112001001010002?00?10?110110[01]001?1100?00?0?00100??01??0?00??10100000?11?010??01?10000??0000 ??????010?000??1?1?0??0000110??1???0211????????

Alligatorium ?0??????1?0000?1000010?111??0?100?1????0??00??11??1??1000???????20?1????00101?101?011111000???1?0 0100???????????10??1???????????????????0???????????????????????0??????????????????0??????????????000??[ 01]?1000?00001????????0211??0?????

Eutretauranosuchus delfsi 203????1?10010111000100111?00?0001001110?000?1112011?1010??0?1?0?121204?00001020111???1??0??0? 1?????????000???00?????0?100????110???????0??00???1???0????10?2???001?0??000?1?110?01?0000000???01 0?00???????????????????????0?????? ?????

Goniopholis 203?1211110010111000100111?0010001002?101000?1112011?1010?10?1?021312?4100[01]0[12]02011?1??1?? 0?00?1200?11?000002100010?1101100??101100?000010010001?1???0000110020000011001000011110?01000 00000000010000????01?????0001100?11??0211??0?????

Pholidosaurus decipiens 212?111101??11?11101100111?00100010?211?100001112111?101??10?100?1311?300???2?0???11?1???0??0?? 2?0?????????????????????1????1?110?????0?0010???????????????0?1?0001???10?001?100?010???0??????010? 00????01?????0001100?11??0211??0????? 00?11??021111030111

Dyrosaurus phosphaticus 002??1?101?010?11?00100011?1010011012?10101001112011?1011?10?10101302?3?00??2?000????????0?00? ?????1???????????????????1????????????????0??0???????????????021?0001????0??????00??1?0000000???010?0 0?11?01?????0001100?11??0211??0?????

Arq. Mus. Nac., Rio de Janeiro, v.65, n.4, p.417-459, out./dez.2007 FIRST CRETACEOUS “PROTOSUCHIAN” FROM GONDWANA 457

Sokotosuchus ianwilsoni 2?2???1101??10????001001???101001?012?1??????1112?11?1?11??0???1?1?0?????????01???????????????????? ????????????????????1????????????????0??0????????????????????0?????????????????????????????????00??????? ????????????????????????????

Pelagosaurus typus 202?1111110011020101000000000000[01]100211010000001101111001001?10001200?30000020000110111?0 000001200011101?00???10???????1?1??????0000??010010?0010???00????0001000011201000011000001?0000 000???010000???0?1?0???000?????????????????????

Teleosauridae [02]02?1111110011020100100000000000110021?01000?001101111001011?1?00120003?000?200002101111?0 000?12000101011?0???10??01001101??1011000011000010100?0??0000??10001000011?010?01110001010000 00000000100001?12010000000011000110?0211??0?????

Metriorhynchidae [02]02?12110100111201011000?0000000110021?0?000?001101111001011?1?001200?300010200002101?11?0 000??????0??012?0???100?01001101??1011?000??000010102?0???000????001000011?01000?11000001000000 00???0100001?1?010000000011000110?0211??0?????

Bernissartia fagessi 203??21111??00111000?00111?001000?002?????0001112?11?10100?0?1???1?1??410010102011?1?11??02002 1110110100000??00????????1????1????????0??0?10???01???0????1?12000001????0??????00??10000????0??01 ???????1?1?0???0001100?11????????0?????

Hylaeochampsa vectiana 00???????11???11????1?01???0????0?002?1?1011?????????101??1??1?????10???????????????????????????????? ??????????0???????10????????????0??0??????0????????????2??00?????????????????????????????01?????????1?0 ???000?????????????????????

Gavialis gangeticus 212?121111001111110110111110010001002110101101112011110110101110[01]131003100012000001111110 131112111100?000002110?100100?101??121100?00000001000101?1?00001?0?20?00011001000011100001?0 000000000010000001001001000001100?11??021111030111

Borealosuchus formidabilis 203?1211110010111000100111?001000100211010111111211111010010?110?1310031000110?0111111111131 11?110?00?000002110?100100?101??11110??000000010001?1???0000110?20?00011001000011100001000000 00000010000001001001000001100?11??021111030111

Crocodylus niloticus 203012111100[01]011100010211110010001002110?01111112011110100101110[01]1310031000100101211111 10131112021100?0000021100100100?101??121100?0000000100110101100001?0?20000011001000011100001 00000000000010000001001001000001100?11??021111030111

Alligator mississippiensis 203112?101?0001110001021111001000?002110101111112011110100101110[01]031203100010020121111111 1311120211?0?00000211001001001101??111000?00000001001[12]01011000011[01]1200000110010000111000 01000000000000100000010010010000011

Arq. Mus. Nac., Rio de Janeiro, v.65, n.4, p.417-459, out./dez.2007 458 L.E.FIORELLI & J.O.CALVO

APPENDIX IV

ANATOMICAL ABREVIATIONS

ab: anterior blade; ac.r: acromial ridge; AM: insertion of the M. ambiens; as: astragalus; Br: origin for the M. brachialis; ca: caudal vertebrae (1 to 5); cap: capitulum; CB: insertion of the M. coracobrachialis brevis; ce: cervical vertebrae (4 to 9); DC: insertion of the M. deltoideus clavicularis; dci: dorsal crest of ilium; di: diapophysis; dpc: deltopectoral crest; do: dorsal vertebrae (1 to 4); f: femur; ff: fossa flexoria; FT: insertion of the M. femorotibialis; FTE: insertion of the M. flexortibialis externus; FTI: insertion of the M. flexortibialis internus; gc: glenoid cavity; GI: origin for the M. gastrocnemius internus; h: humerus; hk: scapular hook; hy: hypapophysis; i: ilium (= il); ip: ischiadic peduncule; is: ischium; it: inner tuber; IT: insertion of the M. iliotibialis; k: keel; lc: lateral condyle; ldr3: third left dorsal rib; lh: left humerus (= lu); lpe: lateroproximal expansion of humerus; lr: left ribs; ls: left scapula; lt2: left tibia (second individual); lf2: left fibula (second individual); mc: medial condyle; mcp.ti: medial condyle process of the femur in the tibia; ne: neural spine; ol: olecranon process; os?: osteoderm?; P: insertion of the M. pectoralis; pa: parapophysis; pap: postacetabular process; pb: posterior blade; pdp: postdiapophyseal process; pp: postparapophyseal process; pr: prezygapophysis; pz: postzygapophysis; pu: pubis; r: radius; ra: radial; rh: right humerus; rf: right femur; ri: right ischium; rp: right pubis; rra: right radius; rs: right scapula; rt.f: right tibia and fibula; rul: right ulna; sa: sacral vertebrae (1 to 2); SC: insertion of the M. scapulocoracoideus; sr: sacral ribs; tp: transverse processes; Tr: origin for the M. triceps brevis; tu: tuberculum; u: ulna; vph: ventroposterior process in caudal vertebrae for hemal arches.

Arq. Mus. Nac., Rio de Janeiro, v.65, n.4, p.417-459, out./dez.2007 FIRST CRETACEOUS “PROTOSUCHIAN” FROM GONDWANA 459

APPENDIX V

Definitions of the nodes used in the text and the phylogenetic results (figure 15) with the diagnoses character of each node. The definitions are based in SERENO et al., 2001 and sensu SERENO et al., 2005:

1 – CROCODYLOMORPHA: The most inclusive clade containng Crocodylus niloticus but not Poposarus gracilis, Gracilisuchus stipanicicorum, chiniquensis, ferratus. 2 – “SPHENOSUCHIA”: The most inclusive clade containing Terrestrisuchus gracilis but not Crocodylus niloticus. Characters 33(1), 105(0), 128(0), 197(1), 220(1). 3 – CROCODYLIFORMES: The least inclusive clade containing Protosuchus richardsoni and Crocodylus niloticus. Characters 1(2), 3(1), 16(1), 24(1), 30(1), 45(1-2), 47(1), 51(1), 65(2), 67(1), 68(1), 80(1), 82(1), 86(1), 95(1), 99(1), 164(1), 166(1), 172(1), 173(1). 4 – PROTOSUCHIA: The most inclusive clade containing Protosuchus richardsoni but not Crocodylus niloticus. Characters 25(0), 55(1), 60(1), 73(2), 140(0), 165(2), 185(1), 215(0). 5 – GOBIOSUCHIDAE: The least inclusive clade containing Gobiosuchus kielanae and Zaraasuchus shepardi. Characters 1(1), 32(1), 75(1), 96(0), 97(1), 174(0), 181(1), 182(1), 183(1), 184(1), 186(1), 187(1), 188(1), 189(1), 190(1), 191(1). 6 – PROTOSUCHIDAE: The least inclusive clade containing Protosuchus richardsoni and Hemiprotosuchus leali. Characters 48(0), 50(1), 74(1), 132(1). 7 – MESOEUCROCODYLIA: The most inclusive clade containing Crocodylus niloticus but not Protosuchus richardsoni. Characters 37(2), 39(1), 41(1), 66(1), 79(1), 84(1), 141(1). 8 – Innominated. Characters 31(1), 113(1), 176(1). 9 – Innominated. Characters 55(1), 143(2), 163(0), 169(1), 178(1). 10 – Innominated. Characters 37(1), 170(1). 11 – Innominated. Characters 91(1); 226 (1). 12 – Neuquensuchus universitas. Characters 173(0), 220(1) 13 – “MESOSUCHIA”: not defined. Characters 10(1), 29(1), 73(2), 119(1), 171(0), 192(1), 197(1), 221(2). 14 – METASUCHIA: The least inclusive clade containing Notosuchus terrestris and Crocodylus niloticus. Characters 15(1), 17(1), 26(1), 67(2), 83(1), 142(0), 167(1). 15 – NEOSUCHIA: The most inclusive clade containing Crocodylus niloticus but not Notosuchus terrestris. Characters 6(1), 29(0), 36(0), 80(0), 140(0), 166(0), 209(0). 16 – EUSUCHIA: The least inclusive clade containing Hylaeochampsa vectiana and Crocodylus niloticus. Characters 43(1), 44(1), 69(0), 71(3), 76(1), 90(1), 91(3), 92(1), 93(1), 110(1), 126(1), 200(0). 17 – PEIROSAURIDAE: The most inclusive clade containing Peirosaurus torminni but not Araripesuchus gomesii, Simosuchus clarki, Notosuchus terrestris, Baurusuchus pachecoi, Crocodylus niloticus. Characters 11(1), 81(1), 105(0), 199(0). 18 – Innominated. Characters 32(1), 74(1), 128(0), 139(0), 140(0). 19 – Innominated (Originally Peirosauridae sensu Gasparini et al., 1991). Characters 120(0). 20 – NOTOSUCHIA: The most inclusive clade containing Notosuchus terrestris but not Crocodylus niloticus. Characters 71(2), 76(1), 90(1), 91(1), 104(2), 123(1), 135(1), 145(0). 21 – SEBECOSUCHIA: No definition has been proposed. Characters 1(1), 3(0), 102(0), 118(1), 120(0), 128(0), 130(1), 156(1), 159(1), 160(1). 22 – Innominated. Characters 9(0), 67(1), 80(0), 156(1), 202(1). 23 – NOTOSUCHIDAE: The most inclusive clade containing Nototsuchus terrestris but not Araripesuchus gomesii, Comahuesuchus brachybuccalis, Simosuchus clarki, Baurusuchus pachecoi, Crocodylus niloticus. Characters 45(2), 105(0), 137(1), 156(0), 176(1), 202(0). 24 – SPHAGESAURIDAE: The most inclusive clade containing Sphagesaurus huenei but not Baurusuchus pachecoi, icaeorhinus, Araripesuchus gomesii, Comahuesuchus brachybuccalis, Simosuchus clarki, Notosuchus terrestris, Crocodylus niloticus. Characters 105(3), 121(1), 124(1).

Arq. Mus. Nac., Rio de Janeiro, v.65, n.4, p.417-459, out./dez.2007

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